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MYCOTAAON

AN INTERNATIONAL JOURNAL DESIGNED TO EXPEDITE PUBLICATION
OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS

WO a WEN ie Gal AE IER AS) ee

COM EP hE ey Neel hoe Bee OU A Rel ERG Yeu UE
GON ofS TN GanO Ea eVini tet Om A G ES

DNC LUD EN Ger ehGUR ES

CO-ED! TORS

G. Ls HENNEBERT
French Language Editor & Book Review Editor

RICHARD P. KORF
English Language Editor & Managing Editor

SUSAN C. GRUFF
Assistant Editor

Published by
MYCOTAXON, LTD., P.O. BOX 264, ITHACA, NY 14850, USA

Printed in the United States of America

TABLE OF CONTENTS, VOLUME THIRTEEN
Nox el April-June 1981

Marcel ewltenGa ln sderemmniscences; RLICHARDEP KORE SD icc ae ck alesse
Dascomycetes Exsiccati, fasc. IV, RICHARD P. KORF & SUSAN C. GRUFF..
Alternarta themes and variations, EMORY G. SIMMONS..................
Studies in tropical Corticiaceae (Basidiomycetes) III. Two new
species of Laxttextum, KURT HJORTSTAM & LEIF RYVARDEN.............
Notes on Zoophthora ocetdentalts (Thaxter) Batko (Entomophthorales:
Entomophthoraceae), RYSZARD MIETKIEWSKI, RICHARD S. SOPER &
DAN LLANE DA LIA ien eee te rece anes ie Wns, ances Meares Lites tts ak sce sai cette coos
New cicada pathogens: Massospora ctcadettae from Australia and
Massospora paharitae from Afghanistan, RICHARD S. SOPER............
beotiaceses Lil. Notes onjselected temperate species referred. Co
Hel Ocrummandenymenoscy Die se KEN De Pe DUMON | eee ane se teseee. eer ws ene ah ass
Lichenes Exsiccati distributed by the University of Colorado Museum,
Boulder: Fascicles 1-15, Nos. 1-600, 1961-1979, WILLIAM A. WEBER..
Cultural studies on Porta etnerascens, P. rivulosa, and P.
subvermtspora (Aphyllophorales, Basidiomycotina), K. K. NAKASONE..
NSLCWESPCCLOSHOLMAMQL Case DAV FU lime ENKING 3 tre cris ens eee seer eS anaes
Candtda paratroptcalts, a new species of Candtda, JOHN G. BAKER,
TRAG PO OALKIN, SDAV IDS He PINCUS? G RICHARDOPS D'AMATO. W236 eee so
Notes on Corticiaceae (Basidiomycetes).
VALIES Twomnewsspecicse cfm 1 crinte ss KURIPHJIORTSTAM.GG .2 oes ts ss
IX. Three new combinations in Hypochntctellum, KURT HJORTSTAM.....
Cladosporium banttanum and its synonym Cladosportum trtchotdes,
MICHAB UA ReEMCGUNNIGSGaDANT Ee BORELL Weems ite ceete canis ett le tm cee. 57s e
A preliminary discomycete flora of Macaronesia:
Part 2, Hyaloscyphaceae subf. Arachnopezizoideae, RICHARD P. KORF.
Part 3, Hyaloscyphaceae subf. Trichoscyphelloideae, LINDA M. KOHN.
Restnomycena gen. nov. (Agaricales), an ally of Aydropus, Mycena
AROMCGeOSUOY A. mo Pensa REDHLADS GRR ROINGER, © net. te i ae Es kis 6
Stnotermttomyces, a new genus of Amanitaceae from Yunnan, China,
MURGANG Semaine to ee eee ect Aree oe retnes tee eterno Tent cr atte ite Se aie vecatare ste Gene a ba
Pypewstuaves imathe: !olyporacede U5, species described Dy) Jren.
Reve tilenmeLE Hah) VAR ENOem rn ea ener Mitte eee cesta eee hele ee este Sat theaters
The publication date,of Arendholz's thesis on leaf-inhabiting
Helotiales, WOLF-RUDIGER: ARENDHOLZ G RICHARD P.SKORE. ie... - oe
An alternative view of certain taxonomic criteria used in the
Entomophthorales (Zygomycetes), RICHARD A. HUMBER........ geeeeeees
Les Hymenochaete a éléments hyméniens pinnatifides, J. C. LEGER.....
Chlortdtum and some other dematiaceous hyphomycetes growing on
decaying wood. Corrections and additions, W. GAMS &@ V.
BHOULBOVA = IEGHOVAR See Gate k tate iene cts eRe Mets mW eeoen urea lencrens ot!
Trichophyton raubttschekit, sp. nov., JULIUS KANE, IRA F. SALKIN,
ERENES WELT ZMANSG UGATHERINERSMLEKAT is sicccte scree coke tere tetetane ote's eons. co) eer tas cate
Cyrenella elegans gen. et sp. nov., a dikaryotic anamorph, S. E.
GOCHEN AUR tatters steels etek ed. re toners ete Gee tet otal eee etre toes, ret MaNe enetn al, cate comets aie tems fa

J.W. CARMICHAEL, W.B. KENDRICK, I.L. CONNERS & Lynne SIGLER; M.
SVRCEK & J. KUBICKA; SCRIPTA MYCOLOGICA no. 9; Martyn J. DIBBEN;
Leif RYVARDEN & Inger JOHANSEN; Takashi MATSUSHIMA; A. NEUHAUS-
LOVA-NOVOTNA & D. GUTHOVA-JARKOVSKA; Helga SCHMITZ; Edmund E.
TYLUTKI; W. GAMS, H.A. VAN DER AA, A.J. VANDER PLAATS-NITERINK,
R.A. SAMSON, & J.A. STALPERS; C.A.N. VAN OORSCHOT; J. JARVA &
E. PARMASTO; Ralph M. SARGENT; G.A. ZENTMYER; Clive JAMES;

ay,
85

105
112

Michael R. MCGINNIS; J.E. SMITH, D.R. BERRY & B. KRISTIANSEN;
Harry J. HUDSON; R.C. COOKE; Adalbert RICKEN; G. BRESADOLA; N.W.
SCHAAD

Notice to subscribers: replacement of missing issueS.........--+-----

NOlez July-September 1981

The genus Pilophorus, H. MARTIN JAHNS..........eeeeec ese ccecrerceeces
New dematiaceous hyphomycetes from tropical rain forest litter,

S. ONOFRI, D. LUNGHINI, A. RAMBELLI & L. LUSTRATI...............-.
Cochliobolus ravenelit sp. nov. and C. tripogonts sp. nov.,

J emclT es ALCORN 9, eaxGeotas. o\ Upto es 20s Woven lo apler Gatto pogeneie era + raceGs ket Ete ueeo ease tee meneueaeee
A psychotropic fungus in Nepal, ROBERT F. SCHROEDER & GASTON GUZMAN.
Ascus structure and function in Cochitobolus species, J. L. ALCORN..
A preliminary discomycete flora of Macaronesia:

Parte oeeGeoglossaceaes RICHARD] Pe KOR aise te reiete lee ele e sdets cle + i ciena ees
The psilopezioid fungi. VII. A new species of Pstlopeata exo

France. DONALD H: PFISTER -G. FRANCOISE CANDOUSSAU one. cis caus os) eee
Notes on soil fungi isolated from a 15-year-old aspen stand in

imnterioryAtlaska wp LOLA Kh. OLIVER SG eKE LTHs VANACLEVE toe oretortesctenees eretets
Coprinus sect. Herbtcolae from Canada, notes on extralimital taxa,

and the taxonomic position of a low temperature basidiomycete

forage crop pathogen from western Canada, S. A. REDHEAD §&

Jie em RAQUA TLR 2. citrate «leila. ctehers Be etorerchnegede > see hsea ec ushel seem teat stctetem tel cae
Selerotinta bresadolae Rick, a taxonomic synonym of Cibortnta

eandoLlecia. (lev. WhetZele (LINDASM ws KOMN tener nis seer tiers ee ore
Halystomyces, a new dematiaceous genus from Arizona's Sonoran

Desertys EMORY AGS eS IMMONS ic neechetuctoiniak oratetesede ersdolets hore! ot cselets ia chy ctseeret ons
Osortomyces, a new genus of the Laboulbeniales from Taiwan,

KAT SOUYURIAGUBRADA te tee eieg. 1st riek. eae sr cokerels fubas olen hate Rers Von ope tocc ots tan emai eae
The genus Codinaea. Three new species from the Americas, ADRIANNA

DREW INGS CGF JregslicgGRANES Sen temgstotzg eh aot. tis Ones sPagins, ve odene teak’ teat ck eee ee
APNeWESPECICS OL, Dacryopiyaratromebraai la Bie LOWS een arene ce one
Notice-slhirds International MycologicalsConeress. ee ee ee

NOLorS October-December 1981

Studies in the lichen genus Psoroma I: Psoroma tenue and Psoroma
ernnamomeia, \ Av HENSSEN®&> Bie RENNERS © cw ale cn, eee een ae eee
An undescribed pleomorphic species of Codinaea, NORIHIRO TOYAZAKI...
The psilopezioid fungi. VIII. Additions to the genus Pachyella,
DONALDOH SS PEISTER = § sFRANCOLSESCANDOUSSAU yr. ure cy cee pete nente. 2 ern
Pithomyees pavgtt, a new combination for Trichocladium pavgtt and
Pithomyces funtculosa, MARY E. PALM & ELWIN L. STEWART............
Validation of the Herpomycetineae and Herpomycetaceae in the
Hbaboulbeniales se SABELLE TT SeTAVARES 2) tears oe ot eee
Erynta (oy somcetes: Entomophthorales): validations and new species.
RICHARD CA SoHUMBER sagen cts ni shrYel val scaled ene eee eet eee ee ee
A review of the nonentomogenous Entomophthorales, BRUCE E. TUCKER...
Erynta (Zygomycetes: Entomophthorales): emendation, synonymy, and
transfers, RICHARD A. HUMBER §& ISRAEL BEN-ZE'EV
Revue des med. G. L. HENNEBERT

2) 4G OS 6. 60) 18: OO) 8 as 0) OO 846) 16.8 (C6. 6: “6: 6: 6 i-«. 6: a: BY igh ie eo) whet e

Brian SUTTON; C.A.N. VAN OORSHOT; K.T. VAN WARMELO & B.C. SUTTON ;
D.M. DRING; J.A. VON ARX; Jan KOHLMEYER & Erika KOHLMEYER; R.C.W.

405

419

A28

431

BERKELEY, G.W. GOODWAY & D.C. ELLWOOD; E.R. WEIBEL; Sultan
AHMAD & S.H. IQBAL; R. VANBREUSEGHEM & Ch. DE VROEY; A. GOFFEAU
& J.M. WIAME; John WEBSTER; M.L. FARR; T.N. LAKHAMPAL & K.G.
MUKERJI; G.M. OLAH, O. REISINGER & G. KILBERTUS; G.M. WATKINS;
John ERIKSSON, Kurt HJORTSTAM & Leif RYVARDEN; Erwin F. SMITH &
Alfred FISCHER; Peitsa MIKOLA; John PEBERDY; R. HARRISON & G.G.

LUNT

Notices

KVEAGT LLCs OC HCTICGSGON PLES S premier ens. tv arm sie ers ise atiet aleta care Sates eee 525

TMCCEMONBNOUSINEE COSTS AN MAD al. We heer ete Rit west male are ot stecaver eres o2o
AU CNOLMLNDE ARE Mercia ea icisnite ois siimte Sek egers coat aeeets starete Mee snet el sas eae stele ate verene : 526
INOUE COMLUNVUSHANG wL UCN MUG Xa vem attests tence erkt te etc Nelc tens oc cusie eel tors: 528
MYCOTAXON@ publication» dates,..12(2)7-sl3 (1b) fand 13(2) 0.0... wees ee 534
REV LOWCIS re ans ere ate cic. e ete tote crete rere at's MPa rae: Mah suet enna etc cela cre witty. 6 555
Errata my O.LUMes.| Ott nese sclera ers lsncy econ sh a et os ge so see tik Ao thn is Baler aruee Patio 535
BEAL AmmeV.O LUMO mE cere tent eteastetee abe terse ome ae eet ore en The! shel onekatal onaiel ohsie ene Hie ee S 555

EVV ata VO lUMe. 1 ON. atte wacle :

I3
]

LIBRARY

ITHACA, N. Y. 14850
@ MAY 1 21981

MYCOT AXON

.«N INTERNATIONAL JOURNAL DESIGN 'O°EXPEDITE PUBLICATION
OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS

Volume XIII April-June 1981 Noel

CONTENTS

Notcerie "Le Galk va TeMINISCENce..0.0...0 eek bs vie yc bat, s ohare s cies RICHARD P. KORF 1
Diecomycetes* Exsiccati ) fase. IV. ss iene cise see RICHARD P. KORF AND SUSAN C. GRUFF be
escriaria themes: and Variations... 27s 4. w'y eke 0% ow eles S dielgieke Bde wee, EMORY G. SIMMONS 16
Studies in tropical Corticiaceae (Basidiomycetes) III. Two new species
ICE LORAIN wie x ata sie hioabelta <a pie Rak awe es KURT HJORTSTAM AND LEIF RYVARDEN 39
Notes on Zoophthora occidentalis (Thaxter) Batko (Entomophthorales:
Entomophthoraceae). RYSZARD MIETKIEWSKI, RICHARD S. SOPER AND STANISLAW BALAZY 4]
New cicada pathogens: Massospora cicadettae from Australia and
Massospora pahartae from Afghanistan...........ccceececceeces RICHARD S. SOPER 50
Leotiaceae III. Notes on selected temperate species referred to Helotiun
EAE Ce PLAREONN etka te Mote batts y's Bin! te. hale oie Peta Binks A¥y Biae § Fok boaters o 6 KENT P. DUMONT 59
Lichenes Exsiccati distributed by the University of Colorado Museum,
Boulder: Fascicles 1-15, Nos. 1-600, 1961-1979...........%... WILLIAM A. WEBER 85
Cultural studies on Porta cinerascens, P. rtvulosa, and P. subvermtspora
Ayn YL opnorales, “Basidiomycobina) 2. o5 eke n> o 24 wes oe siete oh ol K. K. NAKASONE 105
TOM SCE LOS POS MMIC. a Sd cosh Urn Soe. a cig nly dO K viniollals, © uw eho elt gid iwlais DAVID T. JENKINS 112
Candida paratropicalts, a new species of Candida.
JOHN G. BAKER, IRA F. SALKIN, DAVID H. PINCUS AND RICHARD F. D'AMATO 115
Notes on Corticiaceae (Basidiomycetes)
Witt. tiwornew species: of Tubuliorints. Oe .'s iki + Svc wie oly Vite dances | KURT HJORTSTAM 120
IX. Three new combinations in Hypochnictellum.........eeeeveees KURT HJORTSTAM 124
Cladosportum banttanum and its synonym Cladosportum trichotdes.
MICHAEL R. MCGINNIS AND DANTE BORELLI 127
A preliminary discomycete flora of Macaronesia:
Part 2, Hyaloscyphaceae subf. Arachnopezizoideae.............. RICHARD P. KORF 137
Part 3, Hyaloscyphaceae subf. Trichoscyphelloideae.............. LINDA M. KOHN 145
Restnomycena gen. nov. (Agaricales), an ally of Hydropus, Mycena and
BEPC OTNE wahi's Wp che wie PGE ak Vow Fs sina alas cc ahuo ts typ te wceraletara: dicate S. A. REDHEAD AND R. SINGER 150
Stnotermitomyees, a new genus of Amanitaceae from Yunnan, China......... MU ZANG 171
Type studies in the Polyporaceae 13. Species described by J. H. Léveillé.
LEIF RYVARDEN 175
The publication date of Arendholz's thesis on leaf-inhabiting Helotiales.
WOLF-RUDIGER ARENDHOLZ AND RICHARD P. KORF 187
An alternative view of certain taxonomic criteria used in the
Pucomonnthorales? (Zymomycetes) cic to wc das alse le alk bie wishslelele denims RICHARD A. HUMBER 191
Les Hymenochaete a éléments hyméniens pinnatifides.................. J.C. LEGER 241
Chloriditum and some other dematiaceous hyphomycetes growing on decaying
wood, Gorrections. and additions ....5.0 nga + + ite W. GAMS AND V. HOLUBOVA-JECHOVA 257
Trichophyton raubitschekti, sp. nov.

: JULIUS KANE, IRA F. SALKIN, IRENE WEITZMAN AND CATHERINE SMITKA 259
Cyrenella elegans gen. et sp. nov., a dikaryotic anamorph....... S. E. GOCHENAUR 267
NESS LA IES ee SON tae hel ee din a A Saag at oe Sopiat Sam Roos, Dante, a's WedelGie pauns G. L. HENNEBERT 278
Notice to subscribers: replacement of miSSing iSSUES.........ceeeeescecrrcereevees 287

[MYCOTAXON for January-March 1981 (12: 313-548)
was issued March 17, 1981]

ISSN 0093-4666 MYXNAE 13(1) 1-287 (1981)
Library of Congress Catalogue Card Number 74-7903

Published quarterly by MYCOTAXON, Ltd., P.O. Box 264, Ithaca NY 14850
For subscription details, availability in microfilm and microfiche,
and availability of articles as tear sheets, see back cover

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MYCOTAXON

VOIPmeX [Lie Nowele spp al —4 April-June 1981

MARCELLE LE GAL: A REMINISCENCE

RICHARD P. KORF

Plant Pathology Herbarium, Cornell University
Ithaca, New York 14853 USA

Marcelle Le Gal, born Marcelle Choquart on Febru-
ary 14th, 1895 at Amiens, France, died in the same
Cit yarOleilllicr2) aml 9 ame emeiniluencesOonsmycology, sand
particularly on the development of the taxonomy of dis-
comycetes, has been immense, even though she only en-
Teredeilier iieldmwhens shemwase nearly 140 |years of age.
Her early publications dealt primarily with discomycetes
and showed jan) exceptional attention to’ detail and clear
deliniation of characters such that I, young and wholly
inexpernienced,, immediately =accepted her?’ publications as
authoritative and as models of procedure. My own work
One discomycetesm had» sonly sbegtn sin) 1944, 7and) in 1947
Snemecci tm iniecmnermmoOnumentalmeukecnerches sur ales) orne-
mentations sporales des discomycétes operculés," her
just-published doctorate thesis (which was to receive
the French Academy's coveted "Prix Montagne."). I was
Quickiverdrawnmeintop cde lively “correspondence: with “her,
auCusCeSpilcmtlcmciilercicem ile OUlnmages susie ioheeriully
commented upon and examined problem discomycetes that
Deeceh elicrasOVctuaLlcwsVears me nevem Stinting= Ol8 fer time
andsadvicer

lGwas lO49u before we were? to. meet in) person, on
Piemoccaslonmolmdauatripstomburopeans herbaria: when al was
searching for type specimens of species of the -Arachno-
pezizeae for my own thesis studies. At the Laboratoire
de Cryptogamie of the Muséum National d'Histoire Natu-
rellcustine Paris = ilmwas ecordially oreeted; bysProfessor Ro-
ger Heim and his staff, but above all by Mme Le Gal
herself. The memory is one I shall ever cherish, for
she devoted time and energy to showing me her collec-
tions, and the museum's discomycete holdings, inclu-
dingeeivosemo! smerandols (and™ mine), the eminent Emile
Boudier. Quiet-spoken, ever helpful, Marcelle Le Gal
was from that moment on to be a major influence in my
professional life. Her devotion to the study of fungi,

her insight and mastery of the fungi she knew, remain
an inspiration to all who knew her. An unexpected sur-
prise to me as a young graduate student was that she
entertained me with a sumptuous meal in her Paris
apartment, where I was to meet her greatest joy, her
husband, Etienne. They had married in 1922 on her re-
turn from the United States where she had earned her
M.A. at Columbia University.

On several later occasions I was able again to par-
take of Mme Le Gal's hospitality and kindness during
visits to Paris. The last of these was sad beyond any
expectation, for when 1 visited her in 1973 her beloved
husband had only recently died. Her grief had caused
her to withdraw from duties at the museum to her home
in Amiens, and to stop work on the important mono-
graph of the genus Scutellinia that had been occupying
her for over a .decade. When’ she» learned that 1) was to
visit the Paris museum, she made the unprecedented ef-
fort of coming to the museum to talk with me about our
taxonomic problems, and: offered me the loan of “several
of her specimens of a critical genus we discussed s Her
sadness over “the loss' Gof cher) husband wacmepeoiouna,
ands had clearly devastated! herp emotionally imean cerino
both her -personal and, professional mile we siemwacmcoon
to turn her back on mycology, perhaps even on humani-
ty, as the result of an unfortunate accident she suffer-—
ed: in’ Amiens. and) of her grief overmthe Hossmoimherscon—
stant companion-husband. Mycology's great loss is that
her Scutellinia monograph will apparently remain unpub-
lished and incomplete.

Marcelle Le gal is one of the very few to have mar-
kedly affected my development as a scientist. From her
I learned patience and devotion to detailed study of
minute structures as keys to relationships. Not always
was I to be in her good graces; particularly because
of my insistence on following the Code of Nomenclature
and the principle’ of ‘prionity, sl ebecameman tpecemem nad
boy" in her view. For years I licked the wounds inflic-
ted by her vitreolic attack on "le jeune mycologue amé-
ricain" in her masterful diatribe against the Code of
Nomenclature (Le Gal, 1958). That she eventually for-
gave my youthful exuberance was one of my great joys.

I shall miss Marcelle Le Gal's cautionary council
more than many other mycologists will; our interests co-
incided closely, and it is in her footsteps that I have
trod most often, secure in the knowledge that her obser-

MARCELLE LE GAL (1895-1979)

Photographed by the author in the gardens outside the
Muséum National d'Histoire Naturelle, Paris, during
the 1954 International Botanical Congress

vations were dependable and her sense of taxonomic di-
rection unerring. A kinder, more loving spirit has
never before, perhaps, graced mycology.

For those who read French, I commend a touching
tribute to her by Patrick Joly (1980), that captures the
essence of this most wondrous scientist. Yet another
revealing tribute to her contains a complete listing of
her mycological papers, the articlel pymener ) long-time
associate in the field and laboratory, Henri Romagnesi
(1980).

REFERENCES CITED

JOLY, P. 1980. Marcelle Le Gal (1885-1979). Cryptogamie,
Mycol. 1: 93-96.

LE GAL, M. 1958. Petite promenade A travers le maquis de
la nomenclature. Rev. Mycol. (Paris) 23: Lo pee On

ROMAGNESI, H. 1980. Mme Marcelle Le Gal (1895-1979).
Bull. Soc. Mycol. France 96: 125-131.

MYCOTAXON

VOlWexX DL PANO tee pp .seo at April-June 1981

DIS GOMY CETESBEXS NC CAT Ie GRASG® SLV.

RICHARD P. KORF AND SUSAN C. GRUFF

Plant Pathology Herbarium, Cornell University
Ithaca, New York 14853 USA

Twenty-five additional numbers comprise this fourth fascicle
of Discomycetes Exsiccati [for distribution of sets, see the
note on page 15].

We express particular thanks to Dr. Henry Dissing and to Dr.
Sigmund Sivertsen for providing us with holotype material of
Boudtera denntstt Diss. & Siv. in Dissing to issue here as
ISOTYPES (No. 77); a second collection of this species was
collected by the senior author of this paper in their company
and constitutes AUTHENTIC material from a previously unrepor-
Cedmlocali ty, (Now 78),

As previously noted (Dissing §& Korf, 1980), No. 84 in this
fascicle constitutes ISONEOTYPE material of Ruhlandiella bero-
linensts Hennings, the type material of which is presumed to
have been destroyed in Berlin during the Second World War.

An examination of Rehm's type specimen of Awmarta gregarta
Rehm showed that it has warted ascospores, not smooth as is
usually assumed. A smooth-spored variant is issued here (No.
90) as Trtchophaea gregarta (Rehm) Boud. f. laevispora Korf §&
Gruff, f. nov., the specimens issued being ISOTYPES:

Trtchophaeae gregartae f. gregariae similis, sed asco-
sports perfecte laevtbus differt. HOLOTYPUS: CUP-MJ
611 (ISOTYPI itn Korf & Gruff, Dtee. Fxe. 90 disperst).
PARATYPUS: CUP-MJ 41. On clay bank. Ctnehona Botantcal
Garden, elev. 4750 ft., St. Thomas Parish, Jamatca.
Lég. R.P.Korf, leader; J.R.Dtxon, K.P. Dumont, R.W.Erb,
D.H.Pfister, D.R.Reynolds, A.Y.Rossman & G.J.Samuels.
Ciel 71,

The typical, warted-spored form is also issued (No. 89) for
comparison, as Trtchophaea gregarta f. gregarta.

By far the most abundant of Durand's collections of Sarcosoma
eyttartotdes Rehm tm Dur., his No. 1305 (= CUP-A 12278), is

formally designated here as the LECTOTYPE Ofmthaemspecies,
and the specimens issued (No. 79) as Plectanta cyttartotdes
(Rehm in Dur.) Korf are thus ISOLECTOTYPES. [All packets of
this collection in the Durand (CUP-D) and Atkinson (CUP-A)
herbaria in CUP bear the data for collecting site as "Glen
Mary,'' while the original description (Durand, 1903) states,
'Most abundant in Glen Burney.'' This latter name does not
appear on any of the Sarcosoma packets in these herbaria. |

Mr. G. Beaton, of Australia, who has consistently provided

us with exciting discomycete finds, has generously given us
TOPOTYPE material of the elegant species Underwoodia beatontt
Rifai for issue here as No. 91. Dr. Peter Milan Petersen,

of the University of Copenhagen, has contributed the material
from Greenland of Sarecoleotta globosa (Sommerf. : Fr.) Korf
issued as No. 100.

Taxonomic and nomenclatural notes again appear on some of the
labels: (79) on Plectanita cyttartoitdes as the type species of
Plectania sect. Plicosporae; (87) on Karsten's use of the com-
bination Seutellinta seutellata, which while the earliest is
not validly published since the generic name was not yet va-
lidly published; (89) on Karsten's similarly first use of the
combination Sepultarta gregarta, likewise not validly pub-
lished since its generic name had not yet been validly pro-
posed, and on the taxonomic synonymy of Tritchophaea gregarta
var. tntermedia with T. gregarta f. gregaria; (95) on an ab-
berant collection or unnamed variant of Arachnopeztza cornuta
having only 1-celled ascospores; (96) on adoption of Arachnio-
peatza leontna instead of A. candtdo-fulva; (97) on discarding
the group '"'Anomalae" of Arachnopeztza since the "granulations"
on the hairs of A. trabtnellotdes (sole representative of that
group) seem to be more like the resinous excretions sometimes
evident in the group "Typicae" than like the eranulations on
the hairs in Dasyscyphus.

ACKNOWLEDGEMENTS

Particular thanks are due the US National Science Foundation,
which under a series of grants to the senior author has funded
the collecting in Jamaica and the Canary Islands, and to the
Fulbright Commission that supported the collecting in Japan.
The Okinawa collection was by virtue of support from the US
Office of Education, the US Civil Administration of the Ryukyu
Islands, and the University of the Ryukyus. Collections from
Finland and Norway were financed by the University of Copenha-
gen and the University of Trondheim. Dr. William Dress, of
the Bailey Hortorium, Cornell University, kindly checked the
Latin diagnosis.

DISCOMYCETES EXSICCATI

DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

76. Anthracobta macrocystts (Cooke) Boudier, Hist. classif.
discomye. Europes sp...0>, 1907.

On a burnt Eucalyptus sp. clear-cut.

At km mark 14, Bosque de la Esperanza, near Pico de las
Process lenenite. canary iisiands.

beg: R.P.K., W.C.Denison, L.M.Kohn & M.A.Sherwood 8.1.1976
Dele eRe? ake

DISCOMYCETES EXSICCATI

DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

77. Boudtera denntstt Dissing §& Sivertsen tn Dissing, Kew
Bidet eS 5 eel 977 we SOINPE

Near a small stream, Nordland, between Berget and
Fisksjgmoen, 22 km NW of Mo in Rana, Norway.

Leg: H.Dissing, S.Sivertsen §& T.Schumacher GS LX LOS
Deten il. Dissing GvS.Sivertsen

DISCOMYCETES EXSICCATI

DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

78. Boudtera dennisii Dissing §& Sivertsen in Dissing, Kew
Bridle olen owl 97/24 “AUTHENTIC

On mud flat at mouth of brook, among Equtsetum,
Calamagrostts neglecta, and Lamprospora ovalispora.

Loevvaijakka, near Levajok, along Tana River, Finnmark
Fylke, Norway.

Doensosoivertscen,eH sDissing & R.P.K. CARS ARLE ees
Det? Sisivertsen & H.Dissing

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DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

79. Pleetanta cyttartotdes (Rehm tm Durand) Korf, Mycologia
49: 110. 1957. [ISOLECTOTYPE of Sarcosoma eyttartotdes

Rehm 7m Durand]
Oni-deadttwies leaves, etc.

Blue Ridge Mountains, Glen Mary, Blowing Rock, North
Carolina.

NOTES: This is the type species of Plectanita Fuckel sect. Pltcosporae Korf (loc.
@tiajee RUE.K.

Leer eeUrander( oOo) SUL VEIT ELIOT
Deter. TUE

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RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

80) Pseudoplectanta nigrella (Pers. ': Fr.) Fuckel,
Jahrb. Nassauischen Vereins Naturk. 23-24: 324. 1870.

In clay soil on roots of Dtcranopterts linearts and rarely
of Pletoblastus ltnearus.

University of Ryukyus Recreation Area, Oku, Kumigami-son,
Okinawa, Japan.

Roce. ome ete DUNO G mmaekO 1 amar kh sKULOSh ma Heya) Wl bed dS Lore:
Deemer Ke

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RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

81. Ptychoverpa bohemtca (Kromb.) Boudier, Hist. classif.
agScomyc, (Europes) 1 s54eeel O0n.,

On ground in mixed woods.
Lloyd-Cornell Preserve, McLean, New York.

Leg: Mycology Class Zio SS
D6Usa hae ah.

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DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

82. Pulvinula globtfera (Berk. §& Curt. in Berk.) Le Gal,
Prodre, ploresMycolssMadacascar 4-794, 1953,

NUS Oty
otelcOOr Eth elev. in rain) forest, FL NUNGUC we rulert OmR1coOs

Leg: G.Abawi 2ipie L96G
DEUS R TEs Ke

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DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

83. Pulvtnula niveo-alba Moravec, Cesk4 Mykol. 23: 231.
1969.

On duff.
Juuma-Jakalavuoma, Kuusamo, Finland.

Leg: S.Sivertsen Or ES AG
Dette wows.

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DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

84. Ruhlandtella berolinensts Hennings, Hedwigia 42: (24).
1903. ISONEOTYPE, designated by Dissing & Korf,
Mycotaxon 12: 295. 1980.

Oneso bia

Burn site along road in Eucalyptus grove at km mark 14,
Bosque de la Esperanza, Tenerife, Canary Islands.

Legaik.P.K.%, Ro Fogel, GL. Hennebert & LSM /Kohn 299X1T. 1976
pet: HeDussingeg Rey Ke

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DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

85. Seutellinita erinaceus (Pers. : Fr.) O. Kuntze, Revis.
gen. pl. 2: 869. 1891.
On mossy wood.

Michigan Hollow, Danby, New York.

sGamundi ,s Kos. Thind, We. Denison, 9.VIII.1960

Leg: I.J
Ree Moone, | Verelewacorgukers Ks
Ree Kk.

Det:

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DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

86. Seutellinta pennsylvantca (Seaver) Denison, Mycologia
Sila Lobel SO al SOO eae

On old wood.

"Big Woods," SW of Ann Arbor, Michigan.

Leg: L.B.Wehmeyer, R.P.K. G al. 19.VI1I.1948
Det Wc. Denison |

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RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

87. SCcuLeLlinid, scureliata, (Ll. sa Fre) Lambotte., bh leemucols
belpesmoupphs l2299 4 -1837- |

On Fagus sp.

Lloyd-Cornell Preserve, Slaterville, New York.

NOTES: Neither a generic name "Seutellinta (Cooke) Karst." nor the combination

"S. scutellata (L.) Cooke" was validly published > i t
Fl. Fenn. 11: 145. 1884) [ICBN, Arts. 4h, 43]. R.P , arsten (Medd. Soc. Fauna

Leg: M.A.Rosinski Pre1os 2
Det: W.C.Denison

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DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

88. Scutellinta umbrorum (Fr.) Lambotte, Fl. mycol. belge,
Suppl. 1: 300. 1887.

One soil:
Montezuma Wildlife Preserve, Seneca County, New York.

eC cme MUL bed hares 20 GNI LOGZ
DOC aR ek’

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DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

89. Trtchophaea gregaria (Rehm) Boud. f. gregarta, Hist.
Class DiemdiSCONny CmmeEUT Ope map mmo UmLO O74
On soil and rotting plant matter.

Peck Foray, Ashokan, New York.

NOTES: Neither a generic name "Sepultarta (Cooke) Karst." nor the combination
"S. gregarta (Rehm) Karst.'' was validly published by Karsten (Medd. Soc. Fauna
FJ. Fenn. 11: 145. 1884) [ICBN, Arts. 41, 43]. Le Gal's T. gregarta var. inter-
medta Le Gal [Rev. Mycol. (Paris) 2: 214. 1937) is a synonym, since Rehm's type

specimen also has warted ascospores, not smooth as she assumed. R.P.K.
Leg: D.Malloch, R.P.K. & al. TORI ao 2
DeGieR. ake

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DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

90. Trtechophaea gregarta (Rehm) Boud. f. laevtspora Korf §&
GUL ieMy CotaxOnel > omLoo le 1 SULYPE

Onwclayesoilebank:

Trail between Freetown and Wag Water River, near Hardwar
Gap, St. Andrew Parish, Jamaica.

Becca herwe eR aDEx Ole Kab DUNO Coe he Wi 5 Peps INS ea
Dehabrister,. Usk -Reynolds, -A.Y Rossman
§& G.J.Samuels

DOC SRG Pe KersGio Ga.

DISCOMYCETES EXSICCATI

DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

91. Underwoodia beatontt Rifai, Verh. Kon. Ned. Akad.
Wetensch., Afd. Natuurk., Tweede Sect., Su eal Roles.
1968s TOPOTYPE

In sandy soil and debris under Melaleuca lanceolata beside

cliff walk, Anglesea camping ground, Australia.

Leg: G. Beaton ZoeN tile LOGS
Det. Geb.

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DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

92. Urnula cratertum (Schw. : Fr.) Fr., Summa veg. Scand.,
SeCt wy p0S te, p.) 504.8 O40.

On soil and wood.

Woods near Westhaven Road, Town of Ithaca, Tompkins
County, New York.

HEC mn ewe hee OM CN seh el ROLL Gh rake V.1960
Dette RePak.

DISCOMYCETES EXSICCATI

DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

93. Wynnea amertcana Thaxter, Bot. Gaz. 39: 246. 1905.
On soil in woods.

Z-oumiLes-east of Obi, -at crest, of shuld elev 0G te.
Alleghany County, New York.

GeMomith sede biuhms ad. MoROni iG Rar ake ZEN LOO
Deus, Gi GeMio-

ik)

DISCOMYCETES EXSICCATI

DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C., GRUFF, EDITORS

94. Arachnopeaztza aurelta (Pers. : Fr.) Fuckel, Jahrb.
NassaudschenmycreinseNaturk. (2 5224' 1305451870.

On bark and acorns of Quereus sp.
France Brook, Alleghany State Park, New York.

Ibe Oeamhy Det Kean Gala | OAV bOGd
DEC waht Re ke

DISCOMYCETES EXSICCATI

DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

95. Arachnopeztza cornuta Cli Ss aKonieesl boyd 40158"
Tey eae LS) ee (ey ae |

On rotted wood.
Lloyd-Cornell Preserve, Ringwood, New York.

NOTES: Apparently lacking the

2- and 3-septate ascospores by which one usually
recognizes this species. R.P.K.

Leg. Gel-Hennebert (3096) & RYPLK. WV L962
Det raiRe Pike

DISCOMYCETES EXSICCATI

DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

96. Arachnopeztaza leontna (Schw. : Fr.) Dennis, Kew Bull.
Lee OO Leet OOS .

On a very rotted log.
Bast send of Lake Shikotsu, Ibari Pref. , Hokkaido, Japan.
NOTES: This specimen was reported by Korf (Bull. Natl. Sci. Mus. 4: 392, 1959)

as A. eandtdo-fulva (Schw.) Korf. Dennis's transfer is correct, since Schweinitz

aie CL Leontna earlier than Pezisza candtdo-fulva, and Fries accepted

begamoimaigus, Kamei, Y,Otani, R.PvKy 6 al. 20.V.1958
DetewRer Ks

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DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

97. Arachnopeztza trabitnellotdes (Rehm) Korf, Lloydia 14:
POS 9 5 28 ( 8 Goel We

On wood.
Margaretville, Delaware County, New York.

NOTES: The "granulations" on the hairs are more like resinous excretions than
the granulations of Daeyscyphus spp. of the ''Typicae" group. This species be-
longs close to A. cornuta in Arachnopeaiaa group "'Typicae," and should not be

segregated in a group "Anomalae"' as was done by Korf (loc. ctt.). R.P.K.
he Ocarina PelNOY eC taegm habe. eh Gals eye!
DeGeeRn rake

DISCOMYCETES EXSICCATI

DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

O8MParachnopesiza mintopsts /(EI1s Wakore, aMycotaxonmy.
AGS) SESWasy.

Ong bark:
France Brook, Alleghany State Park, New York.

Tee a Rep ko 1G salt Gavi 1o6l
DeGeaR yPaks

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DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

99. Pezoloma larteina (Ell. & Everh.) Korf, Phytologia 21:
2 Ooi Oe. |

On duff, mostly Tsuga needles and cones.
Peck Foray XIV, Twin Valleys, Lewis, New York.

begs Gaapawi, al. PEister 4 Re bak. 15.V1I.1968
Deets RP ak.

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DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY
RICHARD P. KORF & SUSAN C. GRUFF, EDITORS

100. Sarcoleotia globosa (Sommerf.: Fr.) Korf, Phytologia
Ze Orme oe

On Sphagnum fuseum.
Sydpréven (60°21'N, 45 34'W), Greenland.

ever Mialane beter senile OZ) SON LO fel
Deter RP. ke

DISTRIBUTION OF SETS

The first fascicle of this series was deposited in 11 institu-
tions (Korf, 1958), with 2 additional ones listed by Korf and
Gruff (1978) as receiving fascicles II and III. By error they
listed 'BP = Museum of Natural History, Budapest,"' whereas the
sete s actually on depositeat BPUN= Institute of Plant Taxono-
my and Ecology of Edétv6s L. University. A 14th set is now on
deposit at Copenhagen, also beginning with fasc. II. The com-
plete list of deposit herbaria is thus:

BPI (Beltsville) FH (Cambridge) * NY (New York)
BPU (Budapest) * K (Kew) RPG (Paris)

C (Copenhagen) * LAH (Lahore) PR (Prague)
CUP arthaca) MICH (Ann Arbor) UC (Berkeley)
DAOM (Ottawa) UPS (Uppsala)

lack IngmrasciGloml .

REPERENCEs Ci TED

DLSSING gun AND OR cee hORP @LOSOe ePreliminany ssceudres in
the genera Ruhlandiella, Sphaerosoma, and Sphaerozone (or-
der Pezizales). Mycotaxon 12: 287-306.

DURAND, E. J. 1903. The genus Sarcosoma in North America.
Jour. Mycol. 9: 102-104.

ROLE hoes el OooEuea 1 SCOmyCeLeae exSiccatae, asc. 1.) Myco-
LOU a4 Oto oy Been Oona

KORE, oR. P. “AND S.C. GRUFF. 1978. Discomycetes exsiccatti,
Pascal hq ey COtaxXOn a) tl 3o>Z2U0s.

MYCOTAXON

VO XL eNOvae lee p Dram O 704 April-June 1981

ALTERNARIA THEMES AND VARIATIONS
EMORY G. SIMMONS

Department of Botany, University of Massachusetts
Amherst, Massachusetts 01003

PROLOGUS

Being the initiation of a series of observations
on Alternaria species, in this instance on some
whose nonbeaked conidia very rarely occur in
chains (A. molesta sp. nov., A. mouchaccae nom.
nov.) and others whose erostrate conidia prolif-
erate in chains by means of interpolated conidi-
ophores (A. chlamydospora, A. limaciformis sp.
nov., and two commonly misrepresented phytopath-
ogens, A. triticina and A. longipes).

I. EROSTRATAE, SOLITARIAE

Most species of Alternaria produce conidia in chains,
some readily and abundantly under conditions of either
nature or culture (e. g., A. tenuissima (Kunze in Nees &
Nees ex Persoon) Wiltshire), others sparsely and perhaps
only tardily after manipulation of culture medium, light
quality, and temperature. This latter reluctantly catenu-
late group includes, among others, common phytopathogens
whose conidia are thin walled and filamentously rostrate
(e. g., A. zinniae M. B. Ellis) as well as others whose
conidia are relatively thick walled and erostrate (e. g.,
A. radicina Meier, Drechsler & Eddy).

Alternaria species of a third group produce two conid-
ia in a chain so rarely, under any condition reported or
observed thus far, that any chain can be considered little
more than an aberration. Two notable members of this group
are A. chrysanthemi Simmons & Crosier (Simmons 1965) and
A. helianthi (Hansford) Tubaki & Nishihara (1969). Both of
these species are phytopathogens; the nonbeaked conidia of
both are relatively large for the genus, with lengths up to
120um and 110um respectively (Fig. 1).

An additional member of this group with erostrate,
solitary conidia is known in the form of a small-spored
species isolated from a skin lesion of a captive harbour
porpoise (by S. Andersen, Odense, Denmark; transmitted by
G.-A. de Vries, Baarn, The Netherlands, as CBS M 339.77).
It is reported here as novel on the basis of examination of
axenic cultures, primarily on potato-carrot agar, hay
extract agar, and 20% V-8 juice agar (hereafter as PCA, Hay,
V-8; see Stevens 1974), in a diurnal fluorescent light/dark
eyclenpat 22'C™

1. Alternaria molesta Simmons, sp. nov. (Fig. 1)
[molesta = troublesome (to both Phocaena and Simmons,
though in different ways) ]

Ex culturis in agaro PCA descripta. Coloniae griseo-
brunneae, lanuginosae, postea atrobrunneae, applanatae.
Mycelium initio ex hyphis septatis, ramosis, subhyalinis
vel pallide stramineis, levibus, 3.5-5.0um crassis composi-
tum. Conidiophora simplicia, singulatim ex lateribus
hypharum oriunda, recta vel flexuosa, cylindrica, pallide
flavo-brunnea, levia, 0-10 septata, plerumque 4-5um crassa,
usque ad 100 (plerumque 5-20)um longa, apice rotundato et
uniporoso, raro geniculata. Conidia solitaria, subcylin-
drica, ovoidea, obclavata vel ellipsoidea, septis trans-
versalibus plerumque 3-6, longitudinalibus nullis vel
paucis, ad septa definite constricta, pallide straminea,
levia, 15-38 x 7-12 (plerumque 27 x 12)um. Origo typi:
ex laesione pellis Phocaenae phocaenae, Odense, Daniae.
Typus: partes ex Simmons 32-075 (CBS M 339.77) desiccatae
et in BPI, CBS, DAOM, IMI, NY conservandae.

The type isolate is stable and predictable in its
growth and sporulation characteristics under specified
culture conditions. It grows well on PCA and V-8 (15mm
radially in 4 da), producing colonies that are dark olive-
brown with a shallow aerial layer of paler woolly hyphae.
Conidium production occurs initially on hyphae near the
substrate surface but spreads gradually to the intertwined
aerial elements. After the agar surface of an entire
culture plate has been overgrown, the colony becomes almost
Opaque because of the very dark color of both the submerged
and the aerial hyphae. Although the mycelium of colonies
on Hay is scarcely visible, it gives rise to conidia
readily and abundantly.

Conidia of A. molesta are produced one per conidio-
phore tip; conidiophore tip proliferation (geniculation) is
uncommon. Among many hundreds of conidia viewed in slide

preparations, only half-a-dozen had an apical l=cell exten-
sion with a terminal scar, and thus presumably had served
as the basal unit in a chain of two-conidia. The color of
conidium and conidiophore walls is so dilute that septa and
disjunction scars appear in bold contrast in transmitted
light. A very high percentage of conidia mature as ellips-
oid to ovoid units with three transverse septa and at a
size of about 27 x 12um. The conidium population is, for
an Alternaria species, remarkably restricted in morpholog-
ical variation, with very little tendency to produce longi-
tudinally asymmetrical or excessively cellular entities.
Fig. 1 illustrates conidiophores and a representative range
of A. molesta conidium shape, size, and septation. Conid-
jum outlines for A. chrysanthemi (Fig. 1, lower left) and
A. helianthi (Fig. 1, lower right) are included for pur—
poses of comparison.

2. Alternaria mouchaccae (Fig. 2)

Arid and semidesert soils harbor remarkable numbers of
phaeodictyosporic hyphomycete species (Durrell & Shields
1960; Kuehn 1960; Ranzoni 1968; States 1978). New taxa al-
most inevitably have been unearthed and described in Ulo-
cladium, Embellisia, and Alternaria. Pertinent to the
present discussion is a species published as Ulocladium
chlamydosporum Mouchacca (1971), which was described as new
on the basis of several isolates from arid and newly culti-
vated soil of the New Valley region of Egypt. A second |
dematiaceous entity, often associated with U. chlamydospo-
rum in nature and morphologically similar to it, later was
published as a new species Alternaria chlamydospora Mou-
chacca (1973). Study of representative strains.of these
two species (IMI 156434 and IMI 156427 respectively; EGS
31-061 and 31-060) reveals that they are congeneric in
Alternaria., Their, morphologicalwsimilarity andenature
association in the soil tempts one to speculate that they
also may be conspecific although culturally in different
phases of progressive degeneration (a condition noted by
Mouchacca, 1971). However, relying to the greatest extent
possible on comparisons of l-conidium reisolates on PCA and
Hay agars, of conidiogenesis, and of conidium development
before typical cell distortion begins (Figs. 2 & 3), I con-
clude that there are two distinguishable species, Alter-
naria chlamydospora Mouchacca (discussed below as species
no. 3) and Alternaria mouchaccae Simmons, nom. nov. [= Ulo-
cladium chlamydosporum Mouchacca, Revue de Mycologie 36(2):
120, 1971; non A. chlamydospora Mouchacca, 1973].

i oa sal sas
on PCA;
raoh tye

Alternaria molesta ex Type (at top),

chrysanthemi

(Vowerd ihe hoe):

jm @ AL 3k Evioy ie Jo) aL

from cGulLtune

(lower

A. mouchaccae is yet another example of a species that
produces nonbeaked solitary conidia when grown on media of
low nutrient content. False conidial beaks (secondary
conidiophores) are extremely rare under such conditions;
only two were found among hundreds of conidia examined.

A few young conidium initials might be misinterpreted as
ulocladioid on the basis of a narrow isthmian attachment to
the conidiophore. However, most young conidia are spher-
ical or ovoid from the beginning and have bases in close
contact with conidiogenous cells, which themselves exhibit
conspicuously pigmented areas around the terminal disjunc-
tion scars — characters more nearly typical of Alternaria
than of Ulocladium. Conidium basal scars appear always to
have been pushed into an eccentric position following the
division of any basal cell by a vertical or oblique septum.
Enlarging conidia often become conspicuously swollen and
distorted, especially when produced in colonies on V-8 agar.
Thesstrain studied, Vcultura’typica, s(@Mouchacca 1972). aex—
hibits abundant nonsporulating hyphal overgrowth, although
conidium production on nonaerial mycelium remains good as
long as only conidia are used as inoculum in serial
transfers.

Considerable experience with moderately unstable iso-
lates of this sort suggests that hyphal chlamydospores,
stromatic cellular masses, and progressive degeneration
originally described for this material, and still present,
may be characteristic of the typical culture rather than of
the species in toto, and that focusing attention on the |
morphology of conidia before they become distorted may
permit recognition of A. mouchaccae among future isolates,
whether or not accessory structures are present.

II. EROSTRATAE, PROLIFICANTES

The character "conidium with beak," i.,e., a conidium
with an apical portion morphologically distinguishable from
the septate sporebody, strongly influences the commonly
held generic concept of Alternaria. Neergaard (1945) dis-
cussed the nature of the conidial beak, emphasizing its
species-diagnosis utility and making a strong point of
differentiating between "true beak (eurostrum)" as an inte-
gral part of the conidium and "false beak (pseudorostrum),
formed like the conidiophore of the species concerned."

The nature of the Alternaria false beak remains poorly
understood, however, if we judge on the basis of much of
the available descriptive literature and posited ranges of
conidium length. Evaluation of this literature, a complex

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task, is needed in aid of correct Alternaria determina-
tions, especially for pathogens that figure largely and too
often with incorrect names in the international literature.

Conidia of four species discussed in Section I above
typically have no portion that can be distinguished objec-
tively as a beak, true or false (an extremely rare pseudo-
rostrum being exceptional). Conidia of a second, relative-
ly large group of species are characterized as being non-
beaked when produced but having a marked ability to produce
chains through the agency of secondary conidiophores (false
beaks), readily and sometimes even exuberantly.

3. Alternaria chlamydospora Mouchacca (Fig. 3)

The kinds of conidium enlargements that characterize
A. chlamydospora (Mouchacca 1973; his Figs. 2 & 3) could
support a suggestion that mycotaxonomists ignore the fungus
and its name as representing a cultural monstrosity. A high
percentage of conidia become grossly swollen and distorted;
conidiogeny may become so disorganized that lateral hyphal
branches, recognizable as potential conidiophores, some-
times enlarge abruptly terminally into quite recognizable
Alternaria, conidioids + (Mouchacca, cc. ,0110en3 eins bead oF
arising in the tretic mode of conidium ontogeny believed to
be typical of the genus. Most definitions of 'chlamydo-
spore’ would encompass such cellular oddities, even though
some of them, at least, are able to produce typical apical
conidiophores and, presumably, secondary conidia.

Eliminating this organism from taxonomic consideration
would be inappropriate for several reasons, two important
ones being (1) that the conidiogeny apparatus and the
morphology of a respectable portion of conidia are distinc-
tive and predictable up to the point at which individual
cells become aberrant and (2) that recognizable chlamydo-
sporic, conidioid isolates (as contrasted to conidium verum
strains) are not uncommon among isolates of dematiaceous
hyphomycetes and for practical purposes, including medical
and phytopathological, cannot be ignored.

As for A. chlamydospora as a recognizable species, its
individual conidia vera lack a definable beak. However,
apical and even lateribasal secondary conidiophore produc-
tion is common, as is subsequent chain formation. It would
be idle to speculate whether or not this species exists in
nature in a nonchlamydosporic state. Nevertheless, it
would be a fascinating exercise (1) for an experimental
Systematist to attempt to isolate this fungus from whatever
aberrational influence or factor is present and (2) for a

mycopathologist to characterize thes factomeiuscums
4. Alternaria limaciformis (Fig. 4)

A singular isolate received from the Commonwealth
Mycological Institute, Kew, in 1954 has remained nameless
since that time (along with several other such offerings)
while preconceived ideas on Alternaria were being tested
(Simmons 1967, 1978) and somewhat similar genera Ulocladium
and Embellisia (Simmons 1967, 1971) were being segregated,
Conidia of this isolate are solitary for the most part
(well over 90%) and have no definable beak. However, sec-
ondary apical and lateribasal conidiophores are fairly com-
mon, with chains of two conidia equally common and of three
rare but easily detectable in each colony scanned. Conidia
often exhibit one or two minor arcs of curvature, even at
the very young stage preceding septation, thus appearing
somewhat vermiform. As transverse, longitudinal, and ob-
lique septa are formed, especially in the central two quar-
ters of enlarging conidia, the curvature often becomes
accentuated two- or even three-dimensionally. The somewhat
stubby, slightly swollen, sigmoid outline of many conidia
bears a fancied resemblance to that of Limax garden slugs.

Alternaria limaciformis Simmons, sp. nov.

Ex culturis in agaris descripta. Coloniae in PCA
planae, griseo-brunneae, atrantes; in V-8 planae, atro-
brunneae, rubescentes; in Hay inconspicuae attamen conidi-
ogenae. Mycelium ex hyphis dilute brunneis, 3-4um diam.,
necnon atrobrunneae, 6-7"um diam., in funiculis radiantibus
compositum. Conidiophora recta vel acclivia, simplicia vel
ramosa, ex hyphis submersis et aeriis oriunda, usque 5um
diam. et plerumque 150-200um longa, ad quinque genicula
sporifera praebentia. Conidia plerumque solitaria, ali-
quando brevicatenulata; initio ovoidea vel plerumque brevi-
vermicularia, hyalina, levia; denique late ovoidea, ellip-
soidea, flexa vel limaciformia, straminea, 30-45 x 9-18um;
septis transversalibus plerumque tribus ad sex et longitu-
dinalibus obliquisve nullis vel in quoque cellula centrali
uno ad duobus. Habitatio typi: solum prope Goole, Yorks.,
Britannia. Typus: partes ex Simmons 7-086 (IMI 52976, leg.

A. kk. Cottrell 307. 2).desiccatae et in BPE. CBS, DAUM a IML,
NY conservandae.

5. Alternaria triticina (Fig. 5)

A. triticina Prasada & Prabhu (1963), a striking
species originally described from wheat leaves and, for-
tunately, represented in culture by an isolate from the
type material, is another example of an Alternaria with
nonbeaked conidia that readily produce apical outgrowths
and, subsequently, short chains of two and three conidia.
In my experience the great bulk of spotted wheat leaves
that reach laboratories in both tropical and temperate re-
gions as Alternaria specimens bear members of the taper-
beaked Alternaria alternata (Fries) Keissler and A. tenuis-
sima groups and, commonly, the Alternaria state of Pleo-
spora infectoria Fuckel. A. triticina specimens are -so
rare in major Alternaria reference collections, including
my own, that it seems likely that the species itself either
is rare (for such a common and intensively cultivated host)
or that it does not sporulate readily in nature, and thus
escapes discovery on most diseased specimens.

A representative isolate (EGS 17-061, ex ITCCF 1186
G. C.) from the type specimen remains in good sporulating
condition almost 20 years after its isolation. Some of its
distinctive characteristics deserve review, in view of the
circumstance that its original description lacks illustra-
tions and is misleading about conidium surface ornamenta-
tion and that the influential published illustrations of.
M. B. Ellis (1976) appear either to be based on atypical
material or to emphasize features (e. g., conidia gradually
tapering distally and with smooth walls) that are not typi-
cal of the species.

Conidia of typical A. triticina terminate distally in
an obtusely conical cell. Conidia produced in nature and
in culture seldom taper into a rostrate structure, although
the terminal cell often renews growth abruptly and produces
a morphological and functional conidiophore. Conidium size
typically does not exceed 60 x 25um. Ornamentation of the
conidium wall is shallow and inconspicuous, though present,
in the type specimen; in the type isolate verrucae are so
abundant on the conidium surface (6-8 per 10um) and so con-
Spicuous as to make observation of secondary conidial septa
difficult. Young conidia are ovoid, becoming ellipsoid.

A prominent, perhaps useful diagnostic characteristic of
enlarging conidia is the tendency for them to become dis-
tinctly inequilateral.

E
=
Ln

Fionn PWN @euliewuee .

ina ex Type.

JMIPESRMNAVETLA TETea (Le

Pan tee,

6. Alternaria longipes (Fig. 6-8)

The fungus species named Alternaria longipes (Ellis &
Everhart) Mason has suffered considerable indignity at the
hands of mycosystematists, nomenclaturists, phytopatholo-
gists, and tobacco farmers. Amends are due here as an in-
troduction to this further example of a species with beak-
less conidia that readily generate apical conidiophores.

A collection of diseased tobacco leaves was received
bygJcebeeiliis: with avletter=dateds0cts /5 169lye irom
Gerald McCarthy, North Carolina Agricultural Experiment
Station, Raleigh. This material typifies the description
of Macrosporium longipes Ell. & Ev. (1892); it still exists
(NY), packeted in McCarthy's original letter with a scrap
of paper carrying one of Ellis's characteristic sketches of
a conidium and notes: "Brown spot"; "35-45u" (length of
conidium body), "15-30u" (length of basal "stipe," a common
misinterpretation at the time of an apical conidiophore or
beak).

E. W. Mason (1928) transferred the Ellis and Everhart
epithet to Alternaria. His decision to do so was based on
a J. C. F. Hopkins isolate from Rhodesian tobacco of "an
Alternaria of the A. solani type," which large-spored
species the isolate later was considered to be. Mason soon
recognized his misdetermination, inscribing a copy of his
1928 paper sent to John A. Stevenson: "Alas I got the
wrong one. Tisdale is preparing to pounce on me over this."
The error was unfortunate in its influence on phytopatholo-
gists working with tobacco diseases at the time; it was
corrected in the Appendix of Hopkins's Tobacco Diseases
(1956, pp. 163-164), along with comments on three, perhaps
four distinguishable species of Alternaria associated with
tobacco leaf diseases.

W. B. Tisdale and R. F. Wadkins (1931) also proposed a
new combination A. longipes after making a detailed experi-
mental study of the brown spot disease in Florida and of
the fungus considered to be causal. The evidence of their
descriptions and illustrations suggests that they were
working with the same fungus that Ellis and Everhart had
described. The work of Tisdale and Wadkins, because of its
experimental approach, also was influential, but the cor-
rect attribution of the name in Alternaria remains A. lon-
gipes (Ell. & Ev.) Mason by priority of publication.

North Carolina, which supplied the type material of
A. longipes, has continued to support interest and research
in many aspects of this commercially important disease and

associated fungi for about 90 years. Reports on much of
the tobacco brown spot work of this period reached publica-
tion using the name A. longipes for whatever small-spored
species was/were present. The practical difficulty of dis-
tinguishing A. longipes from other species commonly present
on tobacco led G. B. Lucas (1971) to suggest that A. longi-
pes and genericotypical A. alternata are the same species.
The suggestion is seductive in its ease of application to
determinations of field specimens; it is unwarranted, how-
ever, on the basis of the literature on the subject. To the
extent that tobacco disease work of the past decade is
bracketed with the single fungus name A. alternata, the
work requires reconsideration with respect to the identity
of the fungus species actually involved.

Alternaria alternata (the name legitimized at present
over the earlier A. tenuis Nees) may or may not be associ-
ated with brown spot lesions of tobacco in North Carolina;
the species A. longipes, however, certainly has such a re-
lationship with much pertinent material, not only in the
U.S.A. but also in tobacco production areas of other parts
of the world. Many field specimens and isolated strains
among scores that have reached me are readily identifiable
morphologically as A. longipes, and many of the latter have
been characterized to me by the donors as being virulent
pathogens under experimental conditions.

Examples of reputedly virulent strains that are mor-
phologically typical A. longipes include EGS 30-101 (ex
Gaui ewains M-/4.*recd 1973)-"EGS 16-135 (exe Gur a emuuecas
R-66, recd 1963); EGS 30-033 (ex J. R. Stavely A-5, recd
1971); EGS 30-034 (ex J. R. Stavely A-3, recd 1971); EGS
30-080 (ex 1971 Fungicide Oxford #163 (slow), recd 1971);
EGS=1/-1793& 1/-'80" CexeD .3Cy M. Conbett-srecdhoGorvia
IMI 123399 & 123400).

The distinctness of A. longipes from A. alternata and
from the commonly encountered species A. tenuissima is
illustrated (Fig. 6) with outlines of a range of conidia
from the type specimen of each species. An excellent
modern field collection of A. longipes (representing several
received! in 19/3 from BE. K. sobers, slifton,=Georeta ie
EGS 35-0995. 1 lustrated in) Pie av.

In culture A. longipes produces conidia successively
and so rapidly in chains that the transition region between
conidium body and new conidiophore tends to be poorly de-
fined. Nevertheless, most conidia reveal the site of the
change from conidium to beak through a slight outline con-
striction and by reductions in wall ornamentation, color,
and thickness. Conidium illustrations in Fig. 8 emphasize

several features of cultured material: the extraconidial
nature of the conidiophore/beak and the fundamentally
erostrate nature of A. longipes conidia; the somewhat re-
duced size of conidia produced in culture; and the remark-
able thickness of conidium walls, which is observable in
some field material and which is accentuated in conidia
produced in culture on PCA. The comparatively restricted
growth of A. longipes on V-8 agar, stressed by Sobers and
Doupnik (1969) in their studies of the species, has proved
a particularly useful differential diagnostic feature of
pure, stable isolates of the species.

The voluminous international literature and the un-
published virulence/aggressivity information shared with me

50 um z
Pease. Tits AbcCermariagloneipes, representative conidia: fromua

eodern tield collection (Sobers. 19733) EGS 33-095)...

by several phytopathologists suggest that many strains of
morphologically distinct A. longipes are pathogenic to to-
bacco plants, that others have degenerated in pathogenicity
during their years in culture, that other distinguishable
species of Alternaria very commonly are found on and iso-
lated from tobacco lesions, and that the total picture, in-
sofar as species of Alternaria are involved, needs review.

On Ue Ope

Continued support by the several herbaria and mycological
institutions noted by abbreviations in the text, by the
National Science Foundation (DEB79 04189), and by the U. S.
Army Natick Research and Development Command is gratefully
acknowledged.

- OoOo-

LITERATURE CITED

Dunneli=) i W.a8 and, Lb. MosShtields.. 1960 sesFungisusolaued
in culture from soils of the Nevada test site.
Mycologia 52: 636-641.

Bilts, oJ. B.,fand B.. M. Everhart. | 1392.8 iNewsspeci esmon
EUnet. SOUL a MY COLOCY a/sL 3 0=1oD.

Ellis, M. B. 1976. More Dematiaceous Hyphomycetes. Kew.
207. pp.

Hopkins, J..C. KF: 1956... Tobacco Diseases ukew. suliior pp.

Kuehn, H. H. 1960. Fungi of New Mexico. Mycologia
52 JOD L4.,

Lucas, G. B. 1971. Alternaria alternata (Fries) Keissler,
the correct name for A. tenuis and A. longipes. Tobacco
Science sD 5/7 —427

Mason, E. W. 1928. Annotated account of fungi received at
the. Imperial Bureau of Mycology. List» IL) (Fascieleal):.
Kew. 43 pp.

Mouchacca, J. 1971. Une nouvelle espéce du genre Ulo-
cladium Preuss. Revue de Mycol. 36: 114-122.

Mouchacca, J. 1973. Deux Alternaria des sols arides
d'Egypte: <A. chlamydosporum sp. nov. et A. phragmospora
van Emden. Mycopath. Mycol. Appl. 50: 217-225.

Neergaard, P. 1945. Danish Species of Alternaria and
Stemphylium. Copenhagen. 560 + 2 pp.

Prasada, R., and A. S. Prabhu. 1963. Leaf blight of wheat
caused by a new species of Alternaria. Indian Phyto-
path. 15: 292-293.

Ranzoni, F. V. 1968. Fungi isolated in culture from soils
of the Sonoran Desert. Mycologia 60: 356-371.

MmicwS — Alternaria,loneipes conidia: large erostrate from

nature (lower left), smaller pseudorostrate from Hay (right)
and PCA (upper left);arrows indicate conidium/conidiophore
iva mus LteOTnerZ. 0 Nes, Contdium wakls usually ornamented, thick.

Simmons, E. G. 1965. Alternaria chrysanthemi. Mycologia
57: 140-143.

Simmons, E. Gs 1967. Typification of Alteznarga, Sctem-
phylium, and Ulocladium. Mycologia 59: 67-92.

Simmons, E. G. 1971. Helminthosporium allii as type of a
new genus. Mycologia 63: 380-386.

Simmons, E. G. 1978. Alternaria—an exercise in diver=
sity; pp. 130-135, in C. V. Subramanian (ed.), Taxonomy
of Fungi. Madras. 304 pp.

Sobers, E. K.,,.and. 8. Doupnik, Jr. | 1969.5" Morphologye
pathogenicity, and cultural characteristics of single
conidium isolates of Alternaria longipes. Phyto-
pathology 59: 202-205.

States, J. S. 1978. Soil fungi of cool-desert plant com-
munities in Northern Arizona and Southern Utah. Journ.
Arizona-Nevada Acad. Sci. 13: 13-17.

Stevens, R. B. (ed.) 1974. Mycology Guidebook. Seattle.
JO3 pp.

Tisdale, W. B.,-and R.-f.) Wadkins. 1931) “BrownisepoG.on
tobacco caused by Alternaria longipes (E. & E.), n.comb.
Phytopathology 21: 641-660.

Tubaki, K., and N. Nishihara. 1969. Alternaria helianthi
(Hans. Comb. nov. = lrans.eBrit. Mycol se soCemmm~oa:
147-149.

MYCOTAXON

VOU hee NO leap ano 4 0 April-June 1981

STUDIES IN TROPICAL CORTICIACEAE (BASIDIOMYCETES) III.
TWO NEW SPECIES OF LAXITEXTUM

KURT HJORTSTAM
Malaregt. 12, S-44135 Alingsas, Sweden
LEIF RYVARDEN

Botanisk Laboratorium, Univ. of Oslo, P.O. Box 1045
Blindern, Oslo 3, Norway.

SUMMARY
Laxitextum incrustatum nov. sp. and L. lutescens nov. sp.
are described from Africa. A key to the known species in
the genus is provided.

The genus Laxitextum was described by Lentz (1955) ina
rather wide sense, but was later emended and restricted by
BOPdTHe Gl 950) ne Thiesatyperspectesel;.«01COLOrsismwidespread
and is illustrated in Eriksson and Ryvarden (1976).

Pa lbora 105) ereported sis. mbicolOm=sLOMmsoOuthy Arrica,
Nigeria, Kenya and Uganda, while Boidin (1960) reported it
ErOmMesalre. During Cur work = with African corticioid® fungi,
we came across two new species in the genus which are
described below.

Key to species

1. Hymenium and pileus cream to deep yellow, gloeocystidia
dtsolnculvweamy LO1d, eS DOLCS HO, sexu — 5) BUN es sewers Le
lutescens

1. Hymenium white to pale cream, pileus if present, dark
brown, gloeocystidia non-amyloid or only very weakly
SOT SDOLCS Fs —DmUMm LONG cotati eketetces es ceetsl eter arch siete o's 'e6 fete 2

2. Trama and pileus, if present, dark brown, tramal hyphae
browni and- smooth, spores oblong ellipsoid, 4-5 x°2,5-3

Sits amie 6G BitcrcennGer sty Obici os creck OOS L. bicolor
2. Trama cream, pileus not known, tramal hyphae pale yel-

lowish and encrusted, spores broadly ellipsoid 4-5 x

SG a UM Ah OOS ene Sitio Sino Sikes c L. incrustatum

DAXITEXTUM INCRUSTATUM Hijortst. & Ryv. nov. Spec.
Fructificatio mollis, plerumque resupinata, late effusa,
0.2 —- 0.4 mm crassa; tramate distincto evoluto; hymenium
leve, cremeum vel obscure ochraceum, leniter rimosum; sy-
stema hyphale monomiticum; hyphis cum fibulis, rectis, lu-
teobrunneis, reniformibus, plerumque incrustatis, 3.5 - 4
um latis, aliquot oleosis, amyloidibus; gloeocystidia cum
schizopapillis, fusiformia, subulata vel obtusa, 70 - 80 x

5 um, vulgo non-amyloidibus; basidia anguste clavata,

15 =~ 25 x 4 - 5 um, 4 - sterigmatibus; sporis subglobosis
vel ellipsoidibus, echinulatis, (4-)4.5(=5) =x 3-3325 50m,
valde amyloidibus.

HOLOTYPUS: Africa. Tanzania. Arusha Prov. Arusha Nat. park.
Mt. Meru E. slope, road to the crater alt. 1800-2300 m.
Suukebwel973. lo Ryvarden 10108. (0).

PARATYPI: do. L. Ryvarden 10048; Tanzania: Kilimanjaro Prov.
Mt. Kilimanjaro W slope, W. Kilimanjaro Forest Sta. alt.

Ge. 1800 m.110.-11:. Feb.21973 WeRyvarden. 10126; eranga

Prov. Lushoto distr., Usambara Mts. Magamba c. 4 km N of
Lushoto, alt s)1600-2000 meo2 1x22. Feb. 19/32 wry Varden
O75 Oren ene 0.

USA: *hOUuLStana, St..Martinsville, novdate, slegne bemuangs
lois. (TRTC), do. Sept. 32. 1889, Leg. ALB @ banglotsez0y>
(TRIG) Without locality, Sept. 913 ,on, Cake Dvanehesm iy =
inqeon therdround;.C.G.) Llovyan4Gyy ie( TRIG

Fruitbody soft, resupinate and widely effused, but loosen-
ing at the margin which is striate or fibrillose, about
0.2-0.4 mm thick with the trama distinctly developed, hy-
menium smooth, creamish to dull ochraceous, more or less
cracked. Hymenial layer somewhat more dense than in the
other species of the genus. Hyphal system monomitic with
loosely interwoven tramal hyphae, golden yellow, straight
and encrusted. The hyphal wall is thickened and the dia-
meter may vary from 3,5 to 6 um, often with adventitious
septa between each clamped cell. Subhymenial hyphae more
irregular, pale coloured to hyaline. Oleiferous hyphae
occur more or less abundant, usually with a slight amyloid
reaction. All hyphae with clamps. Gloeocystidia arising.
from the tramal layer, fusiform, subulate or with obtuse
apex, sometimes with schizopapilles, generally 70-80 x 5
um, but the length varies considerably and they may some-
times be up to 100 um or more, mostly without amyloid re-
action, negative in benzaldehyde. Basidia narrowly cla-
vate, 15-25 x 4-5 um, with four sterigmata.; Spores) sub=
globose to ellipsoid, echinulate, (4-)4.5(-5) x 3-3.25 um,
strongly amyloid.

Remarks. The species is somewhat similar to L. lutescens
(see below), but is easily distinguished by its slightly
larger spores, encrusted hyphae and paler hymenium.
Furthermore, all known specimens are resupinate. The
hymenial layer is denser and the gloeocystidia are non-
amyloid or only very weakly so. Because of its well deve-
loped subiculum and negative reaction in benzaldehvde

the species seems to belong in Laxitextum and is related
to L. bicolor. However, it also has similarities to a

group of species in eee ee such as G. propinguum
(Jacks.) Parm., G. sibiricum, Parm.,@and Gloeocystidium
lacticolor Bres. (type examined). phe Main differences are
the distinct subiculum in Laxitextum and that oleiferous
hyphae are lacking in these species and the sulfovanilline

reaction sis positive, (except in G. lacticolor?). Another

oi,

Fig. 1. Laxitextum incrustans. a) section through) thes hym=
enium b) basidium c) gloeocystidium d) subicular hyphae
e) spores. From the holotype.

Fig. 2. Laxitextum lutescens. a) section through the fruit-
body b) section through the hymenium c) basidium d) sub-
iculur hyphae e) spores. From the holotype.

species which also comes close to the group is Gloeocysti-
diellum furfuraceum (Bres.) Donk, but it lacks clamps.

We have also compared the new species with Scytinostromella
humifaciens (Burt) Freeman & Petersen because of its some-
what Similar spores and gloeocystidia. However, in that
species the spores are smaller and the gloeocystidia react
positively in benzaldehyde. Furthermore, itke all species

in Scytinostromella, aEsOn pr Humitacvens, Lt 1S) distinctly
dimitic with skeletal hyphae.

Fig.3. Spores of a) Laxitextum bicolor Colles Ryviee 12092) 5)
L. lutescens, holotype c) L. incrustans, holotype d) en-

crusted hyphae from L. incrustans, holotype. SEM by L.
Ryvarden.

LAXITEXTUM LUTESCENS Hjortst. & Ryv. nov. spec.
Fuctificatio resupinata vel distincta reflexa, 0.4-0.6 mm
crassa, supra infuscata; tramate distincto evoluto, 024-
0.5 mm crasso; hymenium leve, cremeum vel lutescens,
leviter rimosum; systema hyphale monomiticum; hyphis cum
fibulis, rectis, levibus, laxe intertextis, luteobrunneis,
circiter 4 um latis; hyphis subhymenialibus irregularibus,
hyalinis; gloeohyphae et gloeocystidia numerosa, amy loides;
basidia’ anguste clavata,,.20=-25. x 7 cum, <4 sterigmatibus;
sporis subglobosis vel ellipsoidibus, echinulatis, 4 x 3-
3.25 um, valde amyloidibus.

HOLOTYPUS: Africa. Ghana. Ashanti Region, Bobiri Forest
Reserve, ab. 30 kmsB Of Kumasi... 16.-22. Apri 6097 4a
Ryvarden 12875 (0).

Fruitbody resupinate to distinctly reflexed, 0.4-0.6 mm
thick, pileus deep yellow to pale brown velutinate, azonate,
trama well developed, pale yellowish brown, about 0.4-0.5
mm thick, hymenium cream-yellowish to strawcoloured with

a light purple-brown tint, slightly cracked. Hyphal sys-
tem monomitic, tramal hyphae loosely interwoven, pale
yellowish brown, 3-5 um wide, smooth and with clamps. Sub-
hymenial hyphae more irregular with the colour pale yellow
or hyaline, mostly collapsed. Oleiferous hyphae abundant,
5-6 um wide, distinctly amyloid and with clamps at long
intervals. Gloeocystidia numerous, arising from the tramal
layer, fusiform, with the apex obtuse, now and then with
schizopapilles, 90-120 x-7-9 um, sometimes up to 150-200

um Or more, distinctly amyloid, but negative in benzalde-
hyde. << Basidia’ narrowly clavate, 20-257 x) 4 umjawatne oun
sterigmata. Spores subglobose to ellipsoid, echinulate,
4X3 S220 UR, eeStrongl ye amyloid.

Remarks. Laxitextum lutescens seems to be well located in
in the genus and is similar to L. bicolor, but delimited by
its deep yellowish colour , the amyloid gloeocystidia and
Oleiferous hyphae and the shorter spores. The tramal
hyphae of L. bicolor are besides more distinctly brown.

References.

Boidin, J. 1958. Heterobasidiomycetes saprophytes et
Homobasidiomycetes resupinés. V. Essai sur le Genre
Stereum Pers. ex S.F. Gray. Rev. Mycol. 23:318-346.

Boidin, J. 1960. Le genre Stereum Pers. s.l.au-iCongo
Belge. Bull. Jard. Bot. Bruxelles 30:51-74,283-355.

Eriksson, J. & Ryvarden, L. 1976. Corticiaceae of North
Europe. 4:549-889.

Lentz, P.L. 1955. Stereum and allied genera of fungi in
the upper Mississippi Valley. U.S. Dept. Agric.
Monogr. 24.

Talbot, P.H.B. 1951. Studies of some South African re-
Supinate Hymenomycetes. Bothalia 6:1-116.

MYCOTAXON

Vole DLE mNOwEl Ss pple41=49 April-June 1981

NOTES ON ZOOPHTHORA OCCIDENTALIS
(THAXTER) BATKO
(ENTOMOPHTHORALES : ENTOMOPHTHORACEAE) |

RYSZARD MIETKIEWSKI

Agricultural and Teachers University
08-110 Siedlce, Poland

RICHARD S. SOPER

USDA Insect Pathology Research Unit
Boyce Thompson Institute
Ithaca, New York 14853

STANISLAW BALAZY

Wielkopolski National Park
62-040 Puszczykowo, Poland

INTRODUCTION

The species Entomophthora occtdentalts (Thaxter)
1888 was described by Thaxter (1888) as a species of
Empusa. He recorded this pathogen as frequently and
commonly infecting aphids on Betula popultfolta Marsh in
Maine, U.S.A. Batko (1964b) renamed this species Zooph-
thord ocetdentalts (Thaxter) and then placed it in the
subgenus Zoophthora Batko (1966b) which includes the group
of species referred to as the "Entomophthora sphaerosperma
group" (Hutchison 1965; Waterhouse 1975; Remaudiere et al
1976; Zimmermann 1978) and recently defined as Zoophthora
Batko sensu stricto (Remaudiére and Hennebert 1980).

ik
This research supported in part by a grant to the
senior author from the International Research Exchange
Board (IREX) New York and by a grant from the U.S.
Department of Agriculture PL-480 Program PL-ARS-/75
(FG-PO-358).

Further notes on the occurence of this species are scarce
and data on its morphology constitute only abbreviated ,
records of the original description. Since the systematics
of species within the subgenus Zoophthora has not yet been
thoroughly defined and because the biology and morpholog-
ical characteristics of Zoophthora phallotdes Batko (1966a)
closely resemble those of Z. occtdentalts, it is useful to
present the results of our observations of these fungi in
recent years.

MATERIALS AND METHODS

In August and September of 1977 in Orono, Maine,
five entomophthoraceous species were discovered on aphids
Myzus persicae Sulz. infesting potatoes: Z. occidentalis,
Erynta neoaphidis (Remaudiére) Batko, Z. exttialts (Hall
and Dunn) Batko, Contdtobolus thrombotdes Drechsler
(= Entomophthora virulenta Hall and Dunn), and Entomoph-
thora major (Thaxter) Gustafsson. Of these, Z. occtden-
talts was predominant. This pathogen was also collected
from Aphts fabae Scopoli on pigweed, Chenopodium album L.
The strains of this species were examined in the laboratory
with application of usual microscopic techniques and suit-
able methods of cultivation on egg yolk and Sabouraud's
maltose agar (EYSMA) and potato-dextrose agar (PDA) in 16
hours photophase. Laboratory infections were also
attempted using several aphid species and caterpillars of
Chortstoneura fumtferana Clem. with conidial showers of
the fungus. On aJuly 13, 19/8; a morphologically -simitar
strain was found in the Wielkopolski National Park, Poland,
on an aphid of the tribe Macrosiphina from Deschampsta
caespitosa.(L.) P.B. in the forest undergrowth. sthese
materials have been preserved as microscopic preparations
in lactophenol with cotton blue (LPCB) and the strains
from the USA also in living cultures on media. In the
course of the studies on the morphology and systematic
position of these strains, a comparison was made with the
holotype of Zoophthora phallotdes, and data on the para-
types of this species were obtained.

RESUETS

A. Morphology and Diagnostics. The infected aphids
produced rhizoids (Fig. 1) from the abdominal part of
their bodies as 1-3 compact bundles which firmly attached
them to plant leaves. The tips of the hyphae forming these
rhizoidal bundles are widened and deeply sinuate (Fig. la).

FIG. 1. Rhizoids of Z. occtdentalts from hosts producing
(aye conidiarvand™(b) resting spores. Bar’=)'(a) 20), (b) 1Op-

During the time of sporulation, the conidiophores covered
the bodies with a compact mass of silver-white to white-
cream mycelium; on the aphid from D. caespttosa the color
of the mycelium was dirty lily. Pseudocystidia tapered
with slightly blunt tips are scarce, and not a constant
feature in the microstructure.

The conidia (Fig. 2) are elongated, with sides bowed
slightly outward; conidia may be slightly curved. The
conidium narrows slightly towards the papilla, and has a
bluntly conical apex. In outline, the papilla is tri-~
angular, and often bears a small central apiculus or arch-
wise and obtuse. The single nucleus is oval or elongate,
and strains well in LPCB; the plasma appears somewhat
hyaline with several granules. Dimensions of the conidia
from aphids or from media differed remarkably: From aphids
they were found to be 25-26:5u X 8-10yn (av. 30 X 9), L/D
2:3-3:5 (av. 3.0). From EYSMA conidia were 24-42y X 7-10,
Cave 318X791) L/D)324—4: 5) (av.2-4.0). Secondary+conidia of
the first order are similar to the primary ones buta little
shorter.

After the transfer of the conidia from EYSMA to PDA,
capillispores are formed in great numbers, and a few
secondary conidia of further orders usually appear; these

PiG. 2 «

elongate and slightly curved. Primary conidia may produce
secondary capillispores at the end of a fine tube (c);
these secondary conidia are more strongly curved than
primary conidia and lack a definite papilla (d). Bar=10p.

Primary® conidial ofl occidentalis (a. bimare

latter conidia are short and considerably different from
the primary, with L/D ratio less than 2. Capillispores
(Fig. 2c,d) are found relatively seldom on aphids. Capil-
lispores from aphids were found to be 20.2-25.1 X 7-9u,
and, from media (EYSMA after transfer to PDA), 22-33.3 X
iJ De esoe5 ey Cave Os7 )

Resting spores (Fig. 3) were formed in artifically
infected aphids or on EYSMA at the temperature 25-27°C.
They are smooth, globose, (20.5) 24-30 (32.5)u diameter,
Witheaslaehteyel low wall 5-6 thicks 90n EYSMA at 25-27°C,
resting spores were larger (20-41.5y) than in the insect.
On the basis of many observations of early starges of spore
development, it is concluded that resting spores are formed
both as azygospores and zygospores, even though the
"classical" types of conjuation noted and illustrated by
Thaxter (1888, p. 171-172) were not observed.

Misidentification of Z. oectdentalts and Z. phallotdes
is possible because of the similarities of their published
spore dimensions and, more importantly because of imprecise
characterizations of the conidial apex in the description
of these species. Thaxter (1888, p. 171) described the
apex of Z. oectdentalts as "tapering strongly" whereas
Batko (1966a, p. 10-11) referred to Z. phallotdes as
"sharply pointed". On the basis of the analysis of differ-
ences described by Remaudiére et al. (1976), studies which
did not consider Z. oectdentalts, the authors intended to
provide a diagnosis for Z. phallotdes. However, the com-
parison of the conidia of both species shows that the
following alternatives would be more suitable:

- conidia cylindrical with parallel
sides in outline, hemispherically
convex papilla and broadly obtuse
SC cm i er ee eer ek ee ee emia LL LOtaeS

- conidia ellipsoidally spindle-
shaped with the sides a little
convex, papilla widely conical
or slightly convex often with
small sharp central apiculus,
top part of the spore tapering
With somewhat blunt apex... ... . 24.. ocetdentdaite

B. Biology and Occurance. JZoophthora occtdentalts
was the most frequent pathogen among entomophthoraceous
fungi collected from the aphid M. perstcae in Orono, Maine,
causing 70-80% among the cases of mycosis; whereas the

FIG. 3. Resting spores of Z. occtdentalts are (a) thin
walled in the early stages of formation and may be ZY Z0-
spores (b); the resting spore contents are granular at
first (c) but contain an oil globule at maturity Cae

Bar = (a-c.d) e720, (bb) Lone.

total mortality caused by entomogenous fungi was very high
at the end of the growing season. On the media used in
this investigation, the fungus grew and developed well, and
produced all forms of spores. Under laboratory conditions,
some species of potato aphids - viz., M. persicae,
Macrostphum euphorbtae Thom., Aphts nasturtit Kalt., and
Acyrtostphon solant Kalt. - were successfully infected with
the spores produced on EYSMA. Attempts at similar infec-
tion against larvae of spruce budworm, C. fumtferana,
failed.

The relative absence of Z. occetdentalts from published
reports probably derived form its misidentification as Z.
radtcans (Brefeld) Batko (= Entomophthora sphaerosperma
Fres.). Even the investigations of Shands et al. (1958;
1962; 1963; 1972) on the entomogenous fungi of potato aphicé
in Maine, carried out between 1952-1972, did not report the
presence of Z. occtdentalts. Likewise, Gustafsson (1965)
does not mention it from Sweden. Petch (1939; 1944) on the
other hand reported it from two localities in Great Britain
but contrary to his usual procedure offered no precise
description. Batko (1962) listed it among Polish
Entomophthoraceae from an aphid on nettle, Urttca dtotca
L., in the Bialowieza National Park, where, despite many
repeated thorough searches, this species was found on few
aphids only in one locality. Thus it may be interesting
to determine the circumstances in which the increase of its
frequency in natural conditions occurs, as well as to
continue attempts to experimental infections in biological
control of aphids.

ACKNOWLEDGEMENTS

The authors wish to give thanks to Dr. Andrzej Batko
(Warsaw University, Department of Plant Systematics and
Geography) for rendering the holotype of Z. phallotdes
accesible, to Dr. Ru%ena Krejzova (Czechoslovak Academy of
Seiences, Laboratory of Insect Pathology, Praha), for
furnishing the data on paratypes of this species, and to
Dr. Danuta Krzywiec (Polish Academy of Sciences, Institute
of Agrobiology Poznan) for identification of the aphids
from Deschampsta caespttosa.

REBERPNCE SSO LED

BATKO, A. 1962. Notes on Entomophthoraceous fungi in
Poland. se encomophnaca, 2: 129-139"

BATKO, A. 1964a. On the new genera: JZoophthora gen. nov.,
Triplospertum (Thaxter) gen. nov. and Entomophaga gen.
nov. (Phycomycetes: Entomophthoraceae). Bull. Acad.
PoleeSci=@ClneLi.. L2C7), Sen Sci ee Lotter azo.

BATKO, A. 1964b. Some new combinations in the fungus
family Entomophthoraceae (Phycomycetes). Bull. Acad.
Pol escis Cle fl.) 12.09), eSer-) Scifi hi0 baaGe OG

BATKO, A. 1966a. A new aphidicolous fungus from Poland
Zoophthora phallotdes sp. nov. (Entomophthoraceae).
Actary Mycol. 2) /—15.

BATKO, A. 1966b. On the subgenera of the fungus genus
Zoophthora Batko 1964 (Entomophthoraceae). Acta
My cole 2: 21521".

GUSTAFFSON, M. 1965. On species of the genus Entomoph-
vnora Fres. in Sweden. I. Classification and
distribution. Lantbrukshoegsk. Ann. 31: 103-212.

HUTCHISON, J.A. 1963. The genus Entomophthora in the
western hemisphere. Trans. Kansas Acad. Sci. 66:
237-254.

PETCH, IT. 1939. Notes 'on‘entomogenous fungi. Trans. Brit.
My COM moOCe m2 O02 ita 27 -— Las.

PETCH, T. 1944. Notes on entomogenous fungi. Trans. Brit.
MyCOl errs OCeag2/Ma Gla 2) Ol—98e

REMAUDIERE G., and G.L. HENNEBERT. 1980. Revision
systematique de Entomophthora aphtdis Hoffm. in
Fres. description de deux nouveaux pathogenes
di aphides. Mycotaxon L269 .5 2 1a

REMAUDIERE , Geo. UKELLER, 9B. PAR LEROK C= Ee LATGE. 1976.
fonmuereciens systematiques et biologiques sur
quelques espéces d'Entomophthora du groupe sphaero-
sperma pathogenes d'insectes (Phycomycetes:
Entomophthoraceae). Entomophaga 21 (2): 163-177.

SHANDS, W.A., S.G. THOMPSON, G.W. SIMPSON, and H.E. WAVE.
1958. Preliminary studies of entomogenous fungi for
control of potato-infesting aphids in Maine. J.
Boon. Ente im Lo4—1 66).

SHANDS, W.A., I.M. HALL, and G.W. SIMPSON. 1962.
Entomophthoraceous fungi attacking the potato aphid
in Northeastern Maine in 1960. J. Econ. Ent. 55:
174-179.

SHANDS, W.A., G.W. SIMPSON, and I.M. HALL. 1963. Impor-
tance of entomogenous fungi in controlling aphids on
potatoes in Northeastern Maine. Bull. Maine Agric.
Expewotis Of +eapDe

SHANDS, W.A., G.W. SIMPSON, I.M. HALL, and C.C. GORDON.
1972. Further evaluation of entomogenous fungi as
a biological control agent of aphid control in
Northeastern Maine. Life Sci. Agric. Maine Exp. Stn.
Tech ebull woo. eeoceDp.

THAXTER, R. 1888. The Entomophthoreae of the United
States. Mem. Boston Soc. Nat. Hist. 4: 133-201.
WATERHOUSE, G.M. 1975. Key to the species Entomophthora

EreS.eDUL em BrLUeMYyCOlmsno0C.1 Oo 4-41.

ZIMMERMANN, G. 1978. Zur Biologie, Untersuchungsmethodik
und Bestimmung von Entomophthoraceen (Phycomycetes:
Entomophthorales) an Blattlausen. Z. angew. Ent.
aan | Jad kara ay

MYCOTAXON

VOU Nee None lem pp.) 00-58 April-June 1981

NEW CICADA PATHOGENS:
MASSOSPORA CICADETTAE FROM AUSTRALIA
AND MASSOSPORA PAHARIAE FROM AFGHANISTAN*

RICHARD S. SOPER

USDA, SHAZAR
Insect Pathology Research Unit
Boyce Thompson Institute at Cornell
Tower Road
Ithaca, New York 14853

INTRODUCTION

The genus Massospora was described by Peck (1879)
and emended by Soper (1974). Forbes (1888) and Thaxter
(1888) working independently, correctly placed this genus
in the Entomophthoraceae. Species of Massospora are
pathogenic to cicadas and thus far are found only in
adults. These fungi attack the reproductive organs of
their host. As the fungus grows, the terminal abdominal
segments of the host slough off revealing either conidia
or resting spores. Both stages do not normally occur in
the same individual. The infected cicadas remain alive
and take an active part in transmission of conidia to
other adult cicadas or in distribution of the resting
spores (Soper et al. 1976a). Only one species, Massos-
pora levtspora Soper, has received extensive epizootio-
logical study (Soper et al. 1976b). From this investi-
gation and the observations of White et al. (1979) on

* Partial support for this project was provided by the

United States/Australian Cooperative Science Program
under travel authorization 2-AS-11 of NSF Contract INT
TR a’ YN OT

Massospora ctcadina Peck, it is apparent that these patho-
gens play an important role in the population dynamics of
their hosts.

Except for one report of Massospora, probably erron-
eously identified as M. ctcaditna, from the Japanese cicada
Platypleura kaempfert F. (Kobayasi 1951), these fungi were
known only in the Western Hemisphere. This paper des-
cribes two new Massospora species which indicates a much
wider distribution by the addition of Afghanistan and
Australian localities. This brings the total of known
Massospora species to 13.

MASSOSPORA CICADETTAE Soper, sp. nov.

CONIDIA ochroleuca in massa, ellipsoideis, papilla
gndistincta, parietibus laevibus, 7.3-9.8 x 20.7-28.1- Um
jo70er 0.8 x 24.7 + 2.1 Um, & +s) binucleata vel inter—
dum trinucleata vel uninucleata, nucleis nunquam bipolari-
bus autem ad partem latissimum conidii locatis. SPORAE
PERDURES pallida vitellina in massa, reticulis profundis
cameras distinctas formantes, cristis angustis laevibus
vel raro minute papillatis, 26.8 - 38.7 Um in diam.

(OG e ae thaes Tenn oe Gane ee-

CONIDIA pale brownish yellow in mass, ellipsoid,
Wetheaneindistinct, papilla, smooth-walled, /.3 = 9.9) x
Bm moe olen (9.0 +05 8ixr 24,7 492, 1 tim, x ts), bi-
nucleate or ocasionally tri- or uninucleate, nuclei
located at the broadest part of the conidium, never bi-
polar. RESTING SPORES pale egg-yolk yellow in mass, with
deep reticulations forming distinct chambers, the narrow
ridges smooth or rarely minutely papillate, 26.8 - 38.7 Lm
Pmeitamn (5 ).2e oo tim, x tis). (Fig. 1)

Holotype: AUSTRALIA: New South Wales: 16 km east of
Hay: Growing in the abdomen of an adult cicada, Crcadetta
murraytensts Distant, Det. M.S. Moulds. Conidial stage.
ieDecembers]979, Coll, M.S) and Bb... Moulds. | CUP:

Paratypes: Same data as holotype. Resting spores
present. CUP. AUSTRALIA: Queensland: Goomeri: near
Kinbombi Falls. Growing in abdomen of Ctcadetta murray-
tensts, Det. M.S. Moulds. Resting spore stage present.
WOeDecember 1976.8 Coll. M.S. and Bus. Moulds. ~ CUP.
Queensland: Aloomba: Growing in abdomen of Cticadetta
puer (Walker), Det. K.J. Chandler. Conidial stage

Figure 1. Spore stages of M. ecteadettae: (A) and (B)
scanning electron micrographs of resting Spores; (C)

resting spore as seen through light microscope; and (D)
conidia.

present. 10 February 1973, Coll. K.J. Chandler. CUP. Tas-
mania: Davenport. Growing in the abdomen of Cicadetta
sp., Conidia and resting spore stages present. No collec-
tion data. South Australia Museum.

Name: The species name of this Massospora is based on
the generic name of its host, Ctcadetta.

Host: Cticadetta murraytensts, Cicadetta puer, and
Citeadetta species (Cicadidae).

This species has the largest conidia yet described
(9.0 x 24.7 um) for Massospora. Although Massospora dor-
tstanae Soper measure 10.8 x 21.8 Um thus approaching
these dimensions, the spore surface is verruculose as
opposed to smooth in M. ctcadettae. The nuclei are dis-
tinctly bipolar in M. dortstanae, and although M. ctca-
dettae is likewise generally binucleate, the nuclei are
positioned centrally. The size and ornamentation of
resting spores (M. ctcadettae) are similar to the Chilean
species Massospora tetttgatts Soper which measures 36.3
Um in diameter. In the absence of conidial characteris-
tics these species can be separated by the general lack
of papillae on the reticulation of M. ctcadettae resting
Spores as compared with numerous minute papillae found
on M. tetttgatts resting spores. The location (Australia)
and host (Crcadetta sp.) are strong circumstantial sup-
port of M. ctcadettae when only resting spores are
present.

The host, C. puer, is a serious pest on sugar cane
in northern Queensland, Australia. Although most cicadas
hnaveavery longslite cycles, viz., 4 to 1/7 years, C. puer
is thought to complete its cycle in one year (K.J. Chand-
ler personal communication). Potentially, M. ctcadettae
could have a correspondingly short life cycle, which
would be the shortest known for any Massospora species.
Intensive epizootiological studies on the periodical
cicadas, Magictcada spp. indicate M. ctcadina does not
occur in the nymphal stages of the host nor in other
genera of cicadas. This indicates M. ctcadina must re-
main in the resting spore state for 17 years between the
synchronized emergence of the adults (Soper et al. 1976a).
A hiatus of 9 years between the occurence of M. levtspora
resting spore production in an isolated population of
Okanagana rimosa (Say) was given as evidence of a corres-
pondingly long survival of M. levtspora (Soper et al.

1976b). These observations strongly suggest that the life
cycles of Massospora species are controlled in some way by
those of their hosts rather than by timing mechanisms
intrinsic to the fungi themselves.

MASSOSPORA PAHARIAEH Soper, sp. nov.

CONIDIA ignotae. SPORAE PERDURES cinnamomeae in
massa, reticulis profundis irregularibus indistinctae,
cameras indistinctas formantes, cristis late separatis
aneustisspapilatais se32.0 — 44 q/epimeiny cL en Oc
2 OM in, Xp S)".

CONIDIA unknown. RESTING SPORES dark red-brown in
mass, with deep, irregular to indistinct reticulations,
ridges widely separated, narrow, bearing numerous minute
papillae s2 68 — 44 e/ Um ane diam .a(39.95 ct 422 6 Um, aceon
Gi Tomer)

Figure 2. Scanning electron micrographs of M. pahartae

resting spores, (A) and (B) note the many papillae on
the reticulations.

Holotype: AFGHANISTAN: Paghman: Growing in the
abdomen of an adult cicada, Paharta casyapae (Distant),
collected on "shade and fruit trees", Det. R. Froeschner.
Resting spores present. 3 July 1963, Coll. E.R. Millet
AF 4 68, 23-14756. CUP.

Name: The species name of this Massospora is based
on the generic name of its host, Faharia.

Host: Homoptera: Paharta casyapae (Cicadidae).

The resting spores of this species are similar to
Massospora levispora Soper in their ornamentation (Soper,
1963, 1974). They can be differentiated by their average
larger size 39.5 Um vs. 34.0 Um and fewer reticulations.
In mass M. levtspora resting spores are yellowish brown
as opposed to the cinnamon brown of M. pahartae.

SNOT Ir Care senO BoRECLES

The following key is a revised version of the origi-
nal (Soper 1974). To identify a species the key can be
entered at any point. As additional characters are exam-
ined, a unique combination will be found for each species.
If only the resting spore is present, it will be necessary
to utilize scanning electron microscopy. Complete des-—
scriptions of species 1-11 are given by Soper (1974).
Species numbers are underlined when the characteristic is
unique.

1. Massospora ctcadina 5. Massospora ocypetes
Peck Soper

2. Massospora sptnosa 6. Massospora tettigatts
Cifera, Machado & Vital Soper’

3. Massospora levtspora 7. Massospora cartnetae
Soper Soper’

4. Massospora dortstanae 8. Massospora diminuta
Soper* Soper

* Several orthographic errors were made in the original
descriptions with respect to Latin endings. These have
since been corrected in the Index of Fungi (1975. 4:312).
The corrected spellings are used here.

9. Massospora platypedtae 2s
Soper

10. Massospora dtceroproctae 13.
Soper*

11. Massospora fidtctnae

Soper

CONIDIAL CH

1-1 Contdtal shape 1-3
aoe UlLipsoidalieZ, 354 339 5
10h, DAs
Doc use form aL
c. globose 7
d. navicular 4
e. obovate 4, 10
ime VOA- Cle m2 du se se Oy ag Lal
g. subglobose 11, 7

1-5
1-2 Number of nuclet

ae One 33en) lye 2

b. two is ue oN 4, Oe le Ue
LOR

Cet hirecw sem 0, el?

d., £our or more 6

RESTING SPORE CH

2-1 Ornamentatton of resting
spore reticulum
absent

a. papillae 2)

b. papillae minute 6

ee opis ae COUNGEe Cine EEO s
dy 16)

de Ppapwulaercruncateg2go.s Ut

2-2 Ridges of rettculum

a. broad forming small chambers

b. irregular forming indistinct

Massospora ctcadettae
Soper

Massospora pahartae
Soper

ARACTERS

Contdtal wall ormamentatton

a. -abSen Gyo yu Ossso mes

De AVGLLUCOSEGs lyse sag,

c. verruculose 4, StL O gael:

1-4 Arrangement of nuclet

ae bipolare2 4 man

random 1, 3, 6, 7, 12

Contdtal length

aa less’ thangs um.

be, 8 to Ol mee een
Tra

oe sLORtowitbe mise eo aera
Cen, ee ely 1G ih

de U5 se tom2 Or tin eee eee
5a 6 saat

@. more, thane2O. lime? sao ae
eh,

ARACTERS

2-3 Resting spore diameter

a. less than 25) Um"3

be eZ Seto eS 0e immo eo st Onmmo
2

Cr 30 ECOR4S) Uimerlt geome mer
Byecaiky Ais 1

d. more than 45 Um 1

chambers 3, 13

narrow forming distinct chambers 1, 2, 5, 8, 11, 12

HOST

3-1 Genus of eteada attacked

Het scarinetdsy. g. Magtetcada 1

De Cicada? <8: h. Okanagana 3

ec. Cteadetta 12 Bh Peak romeem ANS)

d. Dtceroprocta 10 ieee OuU ped ial.

PeeOresrvanay4. —)) k. Quesada 2

hel) peddemenber’ gos Ie 1. Tetttgates 6

DISTRIBUTION

4-1 Collection localttties
Afghanistan 13 Honduras 11
Argentina 5, 7 MeEXTCO e2,pto pL
Australia 12 (ees, SHEE) iy Gy Bs Ee
spel war dh, Phy AN tL 1G) Venezuela 2

ACKNOWLEDGEMENT S

Dr. R. Froeschner, U.S. National Museum, Washington,
DC, Mr. M. Moulds, Sidney, Australia, and Mr. K. Chandler,
Cairns, Australia, provided the identification of the
cicadas. Dr. R. Humber, USDA, SEA-AR Insect Pathology
Research Unit, Ithaca, New York, provided the Latin trans-
lation of the species descriptions and constructive criti-~
cism of the manuscript. Ms. Myra Libott, Cornell Univer-
sity, Ithaca, provided technical support for SEM photo-
micrography. Their help is greatly appreciated.

LITERATURE CITED

Forbes, S.A. 1888. On the present state of our knowledge
concerning insect disease. Psyche 5: 3.

Kobaysai.e)4-01951'. Notes of fungi 2 (1) 9on the newly found
genus Massospora from Japan. J. Jap. Bot. 26: 21-23.

Peck, D-H. 1879. Report of the botanist. NY State Mus.
Nat. History 3lst Ann. Rept. pp. 19-44.

Soper, R.S. 1963. Massospora levispora, a new species of
fungus pathogenic to the cicada, Okanagana rimosa.
Canepn lev bot wae ee oi =/ O-

Soper, R.S. 1974. The genus Massospora entomopathogenic
for cicadas. Part I. Taxonomy of the genus

Massospora. Mycotaxon 1: 13-40.

SOper whe oe eA wae DeLyzer, and) l,f Re) omttlogets ,Oa mec
genus Massospora entomopathogenic for cicadas. Part
II. Biology of Massospora levispora and its host
Okanagana rimosa with notes on Massospora cteadtina
on the periodical cicada. Ann. Entomol. Soc. Am.
(0958 60-957

SOper. Rh OeR Leh oh momitlinw and Awd. DeLyZer ess La AOD.
Epizootiology of Massospora levtspora in an isolated
population of Okanagana rimosa. Ann. Entomol. Soc.
Am. 69: 275-283.

Thaxter, R. 1988. The Entomophthoreae of the United
States. Memoirs Boston Soc. Nat. Hist. 4: 133-201.

White ad. Mi Lloyd sand. H.62ar.e (19/9 Pacihoveeatoc
sion in crowded suburban periodical cicadas: Popu-
Patlonussout otecontrol. BEcoltogygoU-mus05— 315.

MYCOTAXON

VOISE ALIA NOY Lee pp. 59-84 April-June 1981

LEOTIACEAE I11. NOTES ON SELECTED TEMPERATE SPECIES
REFERRED TO HELOTIUM AND HYMENOSCYPHUS

KENT P. DUMONT
The New York Botanical Garden, Bronx, New York 10458

SUMMARY

Fourteen temperate species of Helottum and Hymenoscyphus
were studied and their taxonomic placement discussed. The
accepted species are redescribed and illustrated. .Of the
fourteen, six were shown to be good species of Aymenoscyphus,
four were demonstrated to be taxonomic synonyms of other spe-
cies of Hymenoscyphus, one additional species is a probable
synonym, and three were shown to be members of the Sclerotini-
aceae.

As was recently pointed out by Carpenter (4981), and by Du-
montrGl980: 1981) ithe identification of the majority of the
mMeOtnopicainspecies Ot inoperculateyDiscomycetesmis ditiivcult
Bic matucimes., frustrating dues tosthe lackijof, comprehensive
and updated monographs and floristic studies. Dumont (1980)
pointed out, in his study of Helottum rufo-corneum, that even
Biewmost common species are still datficult to name conmectly
because of the lack of adequate literature descriptions and
because of existing, confusing synonymies.

We have been attempting to work out the names of some of
the more commonly encountered species of Inoperculate Discomy-
meces, witch we have collected (Carpenter G Dumont, 19738;. Car-
penter, 1981; Dumont § Carpenter, 1981; Dumont, 1980; and
Haines, 1980) from neotropical regions. Dumont § Carpenter
(1981) reported the difficulty they encountered while attempt-
ing to identify their own field collections of the Helottum-
Hymenoscyphus complex and Helottum-like organisms from Colom-
bia and adjacent regions and pointed out the.need sto ireinves-=
tigate all of the species of that genus described from the
neotropics. Dumont (1981) presented a preliminary summary of
all of the species of Helottum-Hymenoscyphus reported from the
neotropics. In that study many species were actually excluded
from the genus, placed into synonymy, or redistributed to gen-
Pramin the, Sclerotiniaceae.

While attempting to work out some of the complicated synon-
ymies, such as found in Helottium rufo-corneum, it became nec-
essary to study the literature and specimens of several spe-

cies of temperate members of Helottum-Hymenoscyphus. The pur-
posewin the paper aise to present somer or thesintormacionerc.
cently uncovered while studying the types of several temperate
species described originally as Helotium or placed in Helo-
ttum, in hopes that this will add to the growing information
on the genus Hymenoscyphus.

AS. 1 ebecome more, familian wWithvand study saddi tional speeres
of Helottum-Hymenoscyphus, I am beginning to see the emergence
Of tWORGTS tinct oY OUPINGSLOF SPeCles MIN tie) Cen sr game Ne mmencts
fer to as the Hymenoscyphus caudatus group (which includes the
type species H. fructigenus) and the second the Hymenoseyphus
epupnyylus vroup. in the first group, the apotheoe tamarems tip
itate, and the “sterile tissue (outer ectal excipulum) from the
base-=of the stipe to the margin 1s composed ofva weligduevcl-
oped and well defined textura porrecta to textura prismatica.
Inethe: sf. epvonytlus croup, the apothecia are generaliyvetuyp ius
nates to SsuDStinitate, andl the, outermost» trssuc sor ethemetlpe
and frequently the lower portion of the receptacle adjoining
the stipe as composed of a well to poorly -detined textura
elobulosav to texturasanctlaris.) elt assprobebl es thauecthnesemtwe
distinct estructura leditsmerencesaarcewor tiy -Oferecognl G10 9c oe
ther a separate genus for the #. epiphyliuse group, Or am an-
frageneric, rank might .\bevapplicable. Since, the majorityeort
the species of Helottum-Hymenoscyphus still remain to be exam-
inedyand redescribed, 2t would be premature to, makesaddattrona
modi caLvOnsSs auetnis mOOi1n Gt.

Thesaccompanyinge taxonomic key 1ncludessnote only thnesspce
Giesmstudiedsin tthe present work’ butpencorporaves sa limormerlic
species which) have previously studired(DumonteGy Carpenter,
1981; and Dumont, 1980) and accépt-as valid species of kymeno—
scypnus. « vnerkey shouldinotebesthoughb or as @etini tue
but one to which additional "species should bpemadded atvers they
are studied. The methods used are the same as those reported
bye Dumon tartans. 1.

Key to species of Hymenoscyphus studied

1. Apothecia subsessile, substipitate to turbinate; stipe (or substipe)
and base of receptacle of apothecium to the outside composed of tex-

tura anoularisvor ¢lobulosas =e 2). (Hymenoseyphus eptphyllus group)2.

2. Ascospores 28-42x3-5um, fusoid, anterior end pointed, posterior
ends aErenuiaredy ones nt Moy. seat. eae ee ee eee H. dearnesstt 7p aoc.

eee ASCOSPOLeSH lesSe thane 2oningel ON 0 9 wee eae re anes ne ee eee re

3. Ascospores (7-)8-10(-12)x2.5-3.5(-4.5)um, trapezoidal, obo-
void, generally equilateral, if inequilateral then not flat-

tened on Ones suri ace we. cin .esere eee ener H. immttabults pe 73i

3. Ascospores more than el 2m slong aa cse cect ath oe. te a eee eee Al
4. Apothecia occurring on wood, strongly umbilicate; asco-

spores (15~) 15-18 (=20)x(3-) 4—-5im)) oe es H. umbtltecatus

(See Dumont, 1980)

4. Apothecia occurring on leaves, turbinate, not umbilicate;

ascospores, 15218x5.5=5imae ee eee H. eptphyllus p. 66.

(See Dumont, 1980)

1. Apothecia stipitate; stipe (or substipe, if present) composed of tex-
tura porrecta to textura prismatica (if any globose cells present in

Stipe or receptacle, use first choice)

oe eve ee

MNES Ta, Oeics tad teat Bp oa (Hymenoscyphus caudatus group)5.

5. Ascospores equilateral or if inequilateral then not flattened,
Peleval | Varuale2 O1Gat MCORODOVOLGLOLODDYL LOT... esc taieete ees 6:
6. Apothecia characteristically tany, less than Jmm in diam, dis-
coid; ascospores (10-)11-14(-15)x(3-)4(-5)um on leaves; tem-
DEVAL Cour teas esha A abbey aBhog dy be ony aye tet dress H. translucens p. 80.
6. Apothecia characteristically large, greater than 2mm in dian,
irregular in outline, umbilicate; ascospores (4-)5-7(-9)x1.5-2
Poe PUT RIOR SE OE ry ei eee Re ee ecg ee eal a H. leucopsts
(See Dumont, 1980)
5. Ascospores strongly inequilateral, a high proportion flattened..7.
7. Ascospores with a large 'nuclear staining area" visible in
phloxine, cotton blue and analine blue dyes, rarely l-septate,
(22-) 26-30(-35)x4-5(-6)um, ascus apex papillate; tropical.....
Bee eal Shag 5 Fe ent ak he Males Va Ma hres UE Ae PRN Rae ey. shisn ne Cost aie H. sclerogenus
(See Dumont & Carpenter, 1981)
WN SCOSPOLCSUW1ILNOUD NUCl Cares tAINING Area oon car alaeis occ ele te-0, din 8.
8. Ascospores regularly septate (more than 50% in any amount)
Spe Meee Re Meh Sie aay Roce ote sao eet hats 16 ie het chars ¢ Weatens fel gue ‘ape esgueia lore 9.

9. Ascospores l-septate, (17-)18-22(-24)x4-5(-6)um; apo-
checidewitnouc nairseat, thesbase, ol thesstape. i.e...
ee eee ict ML SOs a) OE ce aoe dab. Cie ayelaces H. mustcola

(See Dumont, 1980)

9. Ascospores 3-septate, (20-)24-30(-35)x4-5(-6)um; apo-
Eheclomwltlenairsea tetlcaDascnOPetiicsS OLD .9,1 ero. 4 asi
SLO Pa Say I Reh. 3 cag tS OOO Pg a A. Peedine

(See Dumont & Carpenter, 1981)
SrA S COSPOLRESEASOD CALE tg aha cate aie ci erred site aie resis > igvarnate é 102
10. Ascospores (16-)18-22(-26)x2-3um, with a basal cilium
Eee PO, Le ate OM ee re Ree Foam ohh wt PE SCULILUS
(See Dumont & Carpenter, 1981)
LORASCOSPOTCS WItnOUtl a1 Dasa lCi LaUMies os sine ce ee oe 1%
Vie enSCOSPOLesenoOked Japl Cally rae cas ties crete: Vze
12. Ascospores (16-)18-23(-30)x3-3.5(-4.5)um,
strongly and obviously hooked, tapering
evaduad 1 yetOethe, baser, ates H. serottnus
(See Dumont’ G Carpenter, 1981}
12. Ascospores (14-)16-23(-26)x4-5(-6)um, if
hooked, only slightly and a few per mount,
abruptly pointed at basal end...H. caudatus
(See Dumont & Carpenter, 1981)
TIP ASCOSDOLeS nO tRnOOKeCdn aD LCA LEY si tlre acre el raN
13. Ascospores (11-)12-15(-16)x(2.5-)3-4um,
with internal, oily-resinous material......

(See Dumont, 1980)
13. Ascospores without internal resinous, oily
MATCTIA IN aimee auch sete ence ate ai ate add. 14.
14. Ascospores frequently curved and then
indented, with anterior end pointed;
apothecial margin with a gelatinous
matrix present; ascospores (10-)12-16
(=18)x3-4(-S)pm-,.f. erraticus ps “66.
14. Ascospores if curved, then not indent-
ed, with anterior end rounded; apothe-
cial margin lacking a gelatinous ma-
trix; ascospores (14-)16-23(-26)x4-5

EXsaptin Poe eee Aa Tee My ek a eC Ree Bete: POG ene 6 © H. caudatus
(See Dumont §& Carpenter, 1981)

1. Helottum albopunetatum Peck, Annual Rep. New York State
MUNG Hat oeLe me 4i/e eee O eos

= Hymenoscyphus albopunetatus (Peck) Kuntze, Revis. gen. Pl. 3(3):
40010 91896.

NOTES. Helottum albopunetatum was described from New York
by Peck (1879) as occurring on leaves, and was fully rede-
scribed and illustrated by White (1943), who accépted™=its
placement in Helotium. It was transferred to Hymenoscyphus by
Kuntze (1898) and accepted in that genus by both Dennis (1964) —
and Arendholz (1979). After having studied the type deposited
ine NYS, I conclude that the species is the same (as fyumenocsey—
phus caudatus, and I place it into synonymy and adopt the name
Hymenoseyphus caudatus which has priority. This species fits
well within my concept of #. caudatus as presented in Dumont §&
Garpente rms Lie |

Holotype: U.S.A., New York Adirondack Mts., C. HY Peck (NYS, NY):
2. Helottum caudatum (Karsten) Velenovsky, Monogr. Discom.
BOHen etic UO.e 1954"

Peztza caudata Karsten, Fungi fenn. exs. 547. 1866.
Hymenoscyphus caudatus (Karsten) Dennis, Persoonia 3: 76. 1964.

NOTES. Hetlottum caudatum was originally described by Kar- |
Stenecromea toliicolous collection made 1n,) Panlandacs peersed |
very common temperate and neotropical species. Dennis (1964) jae
Arendholz (1979), and Dumont. & Carpenter (1981) accept’ the |
species in Fymenoscyphus, a decision with which I concur.” Fou
a full description, illustrations, and discussion, see Dumone
& Carpenter (1981).

3. Helottum conocarpt Seaver §& Waterston, Mycologia 34: 517.
1942.

NOTES. Helottum conocarpt was originally described from
Bermuda on leaves. It was accepted in Helottum by White
(1943), but neither Dennis (1964) nor Arendholz (1979) treated
the species. Dumont (1976) has shown the species to be a mem-
ber of the genus Moellerodtseus (Sclerotiniaceae). See Dumont
(1976) Bloretulivdéescription.!) pilustrationsmandedaccusstone

4. Helottum dearnesstt (Ellis §& Everhart) White, Mycologia
ote AR Te FIGS. 612 am

= Phtalea dearnesstt Ellis §& Everhart, Proc. Adad. Nat. Sci. Philadel-
phia 1893: 146. 1894.

= Hymenoseyphus dearnesstt (Ellis §& Everhart) Kuntze, Revis. gen. Pl.
3(3): 485. 1898.

Apothecial morphology — Apothecia minute, scattered or found in small
groups, subsessile, to ca lmm in diam, ca 1mm high, when fresh disc drying
concave, rehydrating convex. Hymenium when fresh at first subolivaceous,

FIG. 1. Hymenoseyphus dearnessit, holotype ex NY, freehand drawings,

x 1,000. A. Median longitudinal section of an apothecium through margin.
B. An ascus and a paraphysis. C. 6 ascospores drawn after discharge from
ascus.

becoming yellow, drying flesh-colored to ochraceous, rehydrating lighter
and yellowish; receptacle generally concolorous with hymenium when fresh,
dry and rehydrated; stipe coloration difficult to detect owing to small
size, but appearing dark brown.

Apothecial anatomy — Asci 8-spored, 90-110x9-12yum, croziers reported
by White to be absent, presence or absence uncertain owing to poor stain-
ing of the asci, but possibly present, long-cylindric to clavate, taper-
ing towards the base and there generally not becoming expanded to form a
small foot, walls ca lym thick, becoming enlarged at the subpapillate to
papillate apex and there 3-4ym thick; pore J+, the pore walls staining in-
tensely blue in Melzer's Reagent. Ascospores 28-42x3-5ym, details ob-
scured in all collections owing to failure of spores to absorb stain ade-

FIG. 2. Hymenoscyphus dearnesstt, holotype ex NY, freehand drawings.

A. Median longitudinal section of an apothecium at approximately juncture
of stipe and receptacle, x 1,000. B. Habit sketch of an apothecium on the@
Substrates xeca 50"

quately, biseriate to irregularly uniseriate, hyaline, smooth, aseptate,
elongate, fusoid, anterior end pointed, posterior end attenuated and

drawn out to a fine point, a cilium reported by White not observed in

these studies, in outline equilateral, frequently sigmoid and curved, gut- |
tules spherical to irregular in outline, irregularly dispersed in the in-
dividual ascospores and generally filling major portion of ascospores and
separated by narrow bands of cytoplasm. Paraphyses equal to or slightly
exceeding the asci, internally hyaline, occasionally branching towards the
base of the asci, septate, filiform, becoming slightly expanded at the
apex and there 2-4ym wide, walls thin, smooth and hyaline, Subhymenium
absent, without a defined zone or differentiation of tissue beneath the
asci. Medullary excipulum poorly developed, in the flanks consisting of
narrow, parallel hyphae 2-3yum wide, in the center of the receptacle the

parallel hyphae becoming torn apart and interwoven, the individual hyphae
hyaline with walls thickened, refractive and smooth. Ectal excipulum: in-
ner ectal excipulum absent. Outer ectal excipulum non-gelatinized, re-
fractive due to thickened walls of hyphae, to ca 40um wide towards the
margin and to ca 60ym towards the stipe, consisting of hyphae originating
in the stipe, and at the juncture of stipe and receptacle extending almost
perpendicularly to the surface of the apothecium, continuing towards the
margin the individual hyphae extending to the surface at gradually lower
angles until at the margin the hyphae parallel to each other and to the
surface of the apothecium, the individual cells brick-shaped, (6-)8-12
(-20)x4-8um, the walls thickened, hyaline and smooth, at the surface of
the apothecium the apically free cells unmodified and remaining appressed
to the surface. Outer covering layer absent. Hairs absent. Margin nar-
row, consisting of hyphae originating in the receptacle and extending
parallel to the surface, as described for the receptacle, but less refrac-
tive. Stipe to the outside composed of a pigmented zone of irregularly
arranged hyphae, in some areas the individual hyphae losing hyphal orien-
tation, and appearing cellular in composition, the individual cells
brick-shaped, globose or angular, lightly to intensely pigmented, the
walls thickened, non-gelatinized and obviously roughened; inside the pig-
mented zone, the hyphae narrow, 2-3um wide, extending parallel to each
other and to the surface of the apothecium and continuing into the medul-
lary excipulum of the receptacle, the individual cells hyaline or pig-
mented light brown, the walls slightly thickened, lightly pigmented or
hyaline and generally smooth. Hairs absent.

Habitat: On old stems of Stetronema ctltatum.

Etymology of the specific epithet: refers to the collector of the ori-
ginal and type specimen.

Holotype: Canada, London, dead stems of Monarda, May 1980, J. Dearness
1713 (FH). See additional notes below.

Tilustrations: White, Farlowia 1: 615, figs. 31-34. 1944.- Dennis, Per-
Boonia 5: 41, fig. 16. 1964.

NOTES. Phialea déarnessti was described by Ellis | Ever-
Marterromea collection made by J. Dearness (1713) in Canada.
As White (1944) has pointed out the type collection was made
in May 1890; and the host was originally described as Monarda,
but Dearness apparently returned to the same locality in June
and collected additional material of the new species and con-
cluded that the host was not Monarda, but Stetronema ciltatum.
To add to the confusion, Dearness gave the new collection the
same number as the original material, but fortunately they
both do represent the same species.

Kuntze (1898) transferred Phtalea dearnessit to Hymenoscy-
phus; White (1944) placed it in Helottum. Dennis (1964) ac-
cepted its placement in Hymenoscyphus, a decision with which I
concur. The species appears to be most clearly related to
other substipitate species in the genus, such as H. eptphyl-
lus. In these species the stipe is composed of hyphae which
do not form a well-structured textura prismatica or porrecta,
as in species such as Hymenoscyphus caudatus and its related
species, but the hyphae become disoriented with the individual
cells losing hyphal orientation and also losing the layered
effect. In H. eptphyllus, H. timmutible and Helottum mtdlan-

dense the outermost tissue of the stipe toward the receptacle

is formed of textura globulosa to textura angularis, Further, |

in these species, the ectal excipulum towards the stipe is
formed of hyphae originating in the interior of the stipe and

extending nearly perpendicularly or at very high angles to the

surface of the apothecium. From the stipe to the margin, the
hyphae approach the surface at lower angles until at the mar-
gin the hyphae are parallel to the surface. In the #. cauda-
tus group the ectal excipulum is formed of hyphae originating
in the stipe and running parallel (or at a very low angle) to

the surface from the stipe to the margin. Hymenoscyphus dear- |

nesstt is separated from these other species by the size of
LoS HasGOSDOTes..

White (1944), in his description of H. dearnessit, men-
tioned that the ascospores produced a delicate cialium 2:-5-3-5
um long. In the portion of the’ type which i -studicd = tiemace
cospores only poorly absorbed the various stains utilized;
anid, for this reason, I was unable oto observe certatuercacurcs
of the anatomy of the species. I did not observe the cilium
described by White, but agree with him that the ascospores do
become obviously attenuated.

5. Helottum teptphytlum (Persoon ex Persoon)) Fr. > oummanves.
Seand Det 500. O49

Peztza eptphylla Persoon, Ann. Bot. (Usteri) 11: 30. 1794.
Peztza eptphylla Persoon, Mycol. Europ. 1: 295. 1822.

Ht out wi

MicheeAcad . S¢i a9 :) Lime 19292

NOTES. Helottum eptphyllum is a very widely distributed
species in temperate regions. As mentioned by Dumont (1981),
the presumed type of Hymenoscyphus eptphyllus deposited at L
contains only sclerotia) of ‘an unidentified fungus 9) UnGileine

situation of the type of the species is resolved, I am follow-@

ing the concept of the species as set forth by Dennis (1956)
and Arendholz (1979), both of whom illustrate the species.
Dumont E1981) shas ftullysdescribedjand) 11 lustratedgautropres.
variant of the species.

6. Helotium erraticum White, Farlowia 1: 606. 1944. FIG. 3.

Hymenoseyphus eptphyllus (Persoon ex Persoon) Rehm ex Kaufmann, Pap.

Apothecial morphology — Apothecia scattered or rarely gregarious, gen-

erally produced from leaf blades, small, stipitate, 0.5-1.0mm in diam, to
ca 1.0mm high, when fresh cupulate in youth, disc flat with age. Hymenium
when fresh white to off-white, drying bright yellow, dull yellow, ochrace-
ous, or rarely reddish; rehydrating translucent, pallid to yellow; margin

generally concolorous with margin when fresh, dry and rehydrated; recepta-

cle when fresh white, drying slightly lighter than hymenium or concolor-
ous, yellow to occasionally reddish, rehydrating as hymenium or slightly

lighter; stipe generally concolorous with the receptacle when fresh, dry
or rehydrated.

Apothecial anatomy — Asci 8-spored, 85-110x8-llym, produced from small —

replicating croziers, broadly cylindric to long cylindric-clavate, taper-
ing to the base and there becoming slightly expanded to form a small foot,
wall to ca lum thick, enlarged to 2-3ym at the rounded to truncate apex;

A )
1A |

LAS

NAN
:

\ A

pore J+ in Melzer's Reagent. Ascospores (10-)12-16(-18)x3-4(-5)um, bi-
seriate above and uniseriate below, biseriate throughout or less commonly
obliquely uniseriate, hyaline, smooth, aseptate or rarely l-septate (and
then apparently in poorly preserved material), oblong, obovoid, ends
rounded to pointed, in outline inequilateral, flattened on one side and
generally curved and then slipper-shaped, generally anterior end slightly
broader than posterior end, guttules present, more or less equal, irregu-
lar in outline, bipolar, filling about half the spore at each end. Para-
physes equal to or rarely slightly exceeding the asci, rarely branched
near the base of the asci, sparingly septate, internally hyaline, f1ili-
form, becoming slightly expanded at the apex and there 2-4ym wide, walls
thin, hyaline and smooth. Subhymenium not well differentiated from the
medullary excipulum, with a perpendicularly oriented zone to ca 40ym in
the center staining more intensely than the remainder of the receptacle
and composed of parallel, short, broad hyphae and croziers, the individual
cells 2-4(-5)um wide, the walls thin, hyaline and smooth, these hyphae
tightly compact and grading into the upper portion of the medullary excip-
ulum. Medullary excipulum poorly to well developed, hyaline, obconical,
non-refractive, consisting in the center of loosely interwoven, branched,
septate hyphae, towards the flank and subhymenium becoming more lightly
compact and parallel to subparallel, the individual hyphae 2-5um broad,
the walls thin, hyaline and smooth. Ectal excipulum not divided into rec-
ognizable layers, to the outside the hyphae more tightly compact and nar-
rower than the hyphae in the center of the excipulum; in this region
forming a well developed textura porrecta to prismatica, the individual
hyphae hyaline, the walls thickened, hyaline and smooth; these hyphae
grading toward and into the narrow hyphae of the medullary excipulun.
Margin at its base constructed as the upper portionon the flanks out
soon becoming characteristically gelatinized, the spaces between the indi-
vidual hyphae 1-2(-3)um, the apical cells unmodified. Stipe composed as
the flanks, the individual hyphae larger than comparable cells in the
flanks. Hairs absent from the margin, flanks, and stipe.

Habitat: On leaf blades of unidentified species, leaves of Amelanchter
sp., Ulmus sp., Hamamelits sp. and Populus sp., and last year's pods of Ro-
binta pseudo-acacta.

Etymology of the specific epithet: relevance to the fungus uncertain.

Holotype: U.S.A., New York, Coy Glen, near Ithaca, on decaying leaves,
3 Oct 1938, H.sHe Whetzel” G.We bk. White sen. “(White herbenos 25472 leanda
3416, one collection apparently divided into two packets or two specimens.
collected on the same day with identical data.)

Paratype specimens: U.S.A., New York, Malloryville, W. of North Bog, on
leaves of Amelanchter sp., Ulmus sp., and Hamamelis sp., 18 Oct 1941,
H. H. Whetzel & Niederhauser s.n. (FH ex CUP 29658); Newfield Gorge, Itha-
ca, last year's pods of Robinta pseudo-acacta, 26 Oct 1941, H. H. Whetzel
& T. Sproston s.n. (FH ex CUP 29666); Canada, Quebec, Duchesnay, County
Portneuf, on leaves of Populus sp., 25 Aug 1938, H. H, Whetzel §& T. Spros-
tones-n se s(hH exCGUP: 27850).

Illustrations: White, Farlowia 1: 607, figs. 14-18. 1944. Arendholz,
Morphologisch-taxonomische Untersuchengen an blattbewohenden Ascomyceten
aus der Ordnung der Helotiales, pl. 15, fig. 4, 1979.

NOTES. Helottum errattcum was described by White (1944)
who indicated that the species was closely related to members
of the Helottum epiphyllum group: H. eptphyllum 4H. immuta-

bile, Hf. carpinacola; and H. mtdlandense. These latter four
share in common one feature which is absent in #. erraticum.
ingaliy foun the sectal excapulum:at the: juncture. of the recep-
tacle and stipe is composed of globose cells to the outside,
with hyphae progressing gradually from high angles to the sur-
Pacem Migc hes LOWeCrepOltLOn stosparallelyat ythe margin: I.-have
found no globose cells in any of the collections studied here
of H. errattcum.

I conclude that #. errattcum is closely related to Hymeno-
seyphus caudatus. In Helottum errattcum the ascospores are
wenerally ‘curved and frequently indented or slipper-shaped,
while in Hymenoscyphus caudatus the ascospores are inequilat-
Cral and generally not curved ‘and when curved, not slipper-
snaped,,) A high proportion of the ascospores in Helotium erra-
ttcum have ascospores with pointed apices, while this feature
UseabscCitein Humenoscypnus caudatue. »1n #, caudatus the mar-
gin 1S composed of thin-walled, brick-shaped cells, while in
Helottum errattcum the margin is unusual for a member of dy-
menoscyphus. The margin is composed of narrow cells apparent-
iMeribeadcodalled Selacinous Matwixn, atestimes, nowever, the hy-
Dpidemalcumientiy compact, and it isediiitecult to; anterpret ‘the
SaMicticeemoncetiec Wa licemerely=appears thickened. ll Ones cole
mectione (CUP sz 9000 lf the hyphae in the ectal excipulum are nar=
rower than normal and very thick-walled, and the ascospores
momitsceDtatcCam Ine collection appears cto ve Dadly preserved.
Waichemaye account 1Orathe abnormal tOrmation of the ‘sterile:
Piscticmangumtnemscptatloleinwtne, ascospores a All ouner tea
Muresmare Cnaracterislics:0n welotrum errartecum,. The shape
a eo ce Se oN Or etic SNCCL1eS. (oS 1S stne Characteristic
Suphymenium i conclude sthat this is merely a’ variant collec-
cron.

I am somewhat concerned about the placement of this species
inp hymenoscyphue, since it produces a margin with at least
some hyphae in a gelatinous matrix. This feature is more
characteristic of the genus Croctcreas than it is of Hymeno-
seyphus, However, the ectal excipulum is that of an Hymeno-
scyphus. Dennis (1964) did not treat the species, while
Arendholz (1979) did place it in Hymenoscyphus as H. erratt-
eus. I conclude that the species is perhaps intermediate be-
tween the two, but shows more affinities to Hymenoscyphus than
Lomcrocicreas. “lowil ly thus; maintain atin Aymenoecyuphust

7. Hetotium fastidtosum Peck, Annual Rep. New York State Mus,
ulate gO 7 eee LiSs/) Sys Fl Gey. 4:

Apothecial morphology — Apothecia scattered, stipitate, 1-1,5mm in
diam, 1-1.5(-3)mm high, when fresh disc flat, drying flat to slightly cu-
pulate, rehydrating convex. Hymenium when fresh off-white to pale yellow,
drying darker, yellow, ochraceous to flesh-colored, rehydrating lighter,
slightly translucent; receptacle when fresh generally slightly darker than
hymenium and white to dark yellow, drying and rehydrating darker than hy-
menium; stipe narrow, cylindrical, when fresh above concolorous with the
lower portion of the receptacle, becoming darker toward the base, drying
darker, especially toward the base, or rehydrating lighter and pallid,
darker below.

Apothecial anatomy — Asci 8-spored, (75-)85-100x9-12um, produced from
small croziers, long cylindric-clavate, tapering gradually to the base and
there occasionally becoming slightly expanded to form a small foot, wall
to ca lum thick, enlarged at the papillate apex and there 2-3ym thick;
pore J+, visible as two, dark blue apical dots in Melzer's Reagent. Asco-
spores (23-)26-30(-35)x2.5-3.5(-4.5)ym, obliquely uniseriate to irregular-
ly arranged, hyaline, smooth, aseptate, subfusoid to clavate, apical end
hooked and pointed, basal end pointed, in outline inequilateral, curved
and rounded in the upper portion, flattened and straight below, anterior
end broader than posterior, with a single row of spherical to irregular
guttules filling majority of the spores. Paraphyses equal to or slightly
exceeding the asci by 5ym, internally hyaline and devoid of pigmented con-
tents, branching occasionally toward the base of the asci, septate, fili-
form, not expanded at the apex and there 2-3ym wide, walls thin, smooth
and hyaline. Subhymenium not well differentiated from the medullary ex-
cipulum, poorly developed, with an indistinct hyaline zone beneath the
asci with hyphae more tightly compact than the medullary excipulum below.
Medullary excipulum poorly developed in the flanks and in the center of
the apothecium, hyaline, non-refractive, consisting of septate, branched,
vertically oriented to tightly interwoven hyphae 3-6um wide, the walls
thin, hyaline and smooth. Ectal excipulum: inner ectal excipulum well de-
fined and well differentiated from the outer ectal excipulum and grading
into the medullary excipulum, entire layer hyaline, non-refractive, to ca
10um wide toward the margin and to 25ym toward the stipe, consisting of
tightly compact, hyaline hyphae 2-4ym wide, with walls thin, non-refrac-
tive, hyaline and smooth. Outer ectal excipulum non-gelatinized, non-re-
fractive or rarely appearing slightly refractive due to thickened walls,
to ca 20um broad toward the margin and to ca 30m toward the stipe, con-
sisting predominantly of a well defined textura prismatica with the indi-
vidual hyphae extending parallel to the surface of the apothecium and
without apically free hyphal tips; the individual cells toward the margin

8-15x4-8um, 12-25x5-8ym toward the stipe, the walls thin or rarely slight- ~

ly thickened, hyaline and smooth. Outer covering layer present or becom-
ing detached and the apothecium then appearing naked, consisting of 1-2
layers 2-5ym broad, the individual hyphae extending parallel to the sur-
face, overlapping, terminating before the margin, the apically free cells
remaining appressed to the surface of the apothecium and unmodified, the
individual cells hyaline or rarely light yellow-brown, the walls non-re-
fractive, hyaline to light brown, and smooth. Hairs absent. Margin gen-
erally poorly developed, narrow above, broader below, entire zone hyaline
or light brown, constructed similarly to the receptacle below, the indi-
vidual cells approximately same size as upper receptacle or slightly
smaller, apical cells unmodified. Stipe in the upper portion constructed
Similarly to the lower portion of the receptacle and apothecial flank, at
approximately midpoint and to the outside 1-2 layers of narrow, hyaline

or lightly pigmented hyphae (often difficult to detect) with thin, lightly
to intensely pigmented and smooth or rarely roughened walls, to the inside
a zone of textura prismatica with the individual cells hyaline to light
brown, in the central core the hyphae narrow, parallel to slightly inter-
woven; hairs absent.

FIG, 4. Hymenoseyphus fasttdtosus, holotype ex NYS, freehand drawings,

x 1,000. A. Median longitudinal section of an apothecium at approximately
midpoint between margin and stipe. B, Median longitudinal section of an
apothecium through margin. C. An ascus with 8 ascospores. D. A branching
paraphysis. E. 7 ascospores drawn after discharge from ascus.

- aN

NAN i} )
NUSag Maes
Nr Ne . an

Ava

Habitat: On leaves of Alnus tneana, Alnus sp., and catkins of Alnus.
Etymology of the specific epithet: relevance to the fungus uncertain.

Holotype: U.S.A., New York, Forestburgh, on fallen petioles of Alnus
leavyes* eocpie, (Ca Hom reckus.D.(NYS),

Additional specimens examined: U.S.A., New York, Adirondack Mts., on
leaves of Alnus sp., date not given, C. H. Peck s.n. (NYS); Labrador Lake,
near Tully, on old overwintered leaves of Alnus ineana, 26 Aug 1935, H. Hy
Whetzel & W. L. White s.n. (CUP 24817 ex FH); Lloyd Preserve, McLean; on
Alnus tncana, leaves on ground, 6 Sept 1935, H. H. Whetzel, W. L. White &
Rogers (White 2042 ex FH). Oregon, Tilly Jane Creek, Mt. Hood, on dead
alder leaves and petioles, 26 Aug 1933, J. Kienholz K 137 (BPI ex FH).
CANADA, Quebec, Duchesnay, County Portneuf, on petioles and lower midribs
of Alnus leaves, 26 Aug 1938, H. H. Whetzel s.n. (FH); locality as previ-
ous. collection, 24 Aug 1958.,.H. HL Whetzely sans (fA):

Tilustrations: White, Mycologia 34: 1158, fig. 2; 166,, fig. (93 1942"
White, Farlowia 1: 153, fig. 7. 1943. Arendholz, Morphologisch-taxono-
mische Untersuchungen an blattbewohenden Ascomyceten aus der Ordnung der
HelOtlales epi peo eld On eee LOO.

NOTES. Helotium fasttditosum was described by Peck from the
Uport.as OCGUTTING Jon, beavesvon Alnus Pe lheuspeciesmuce san om
produce a stroma and is ‘surely referrable to Hymenoscyphus as
reported recently by Arendholz (1979) who also reported it foy
Cheetirsiatimestrom Europe, “Structurally stnessterie surssuc
of the apothecium is similar to the common species Hymenoscy-
phus caudatus and Hymenoscyphus serottnus, while the asco-
spore shape 1S like {that of 4. serozinus = Dumont GeGarnenger
(LOSI) spointed out -thateuntiie recently #.Mecrotiiucewosmecie
erally regarded as /a species Occurring .onehey bac eousme tens
However, they have demonstrated from their studies of neotrop-
ical collections that the species jis extremely variable wand
occurs on leaves as well as on stems. They further reported

the ascospores to be (16-)18-23(-30)x3-3.5(€-5)um and indicatedm
that the base of the stipe generally stains light pink’ in Melg

zer's Reagent,. a reaction also observed in the material stud-
ied here. Dennis (1964) reported the ascospore measurements
for H. serottnus to be 18-28x3-4um, while Seaver reported them
to be 4x22-24um & Rehm (1893 in 1887-1896) found them to be
30-36x4-6uym. The ascospores studied in the present investiga-
tion of Helottum fastidtosum were (23-)26-30x2.5-3.5(-4.5)um
and would thus appear to fall within the reported range for
hymenoscyphus serotinus, and is a probable synonym of #. sero-
tinus., I have been unable to find a-type, of #. Sserottinuc) ang
am thus following the generally accepted concept as presented
by Dennis (1956, 1964). A final decision on the, tentative
Synonymy can be made only after type material is located or a
neotype designated for H. serotinus.

8. Helottum fraternum Peck, Annual Rep. New York State Mus.
Deg) AF 18:79

= Hymenoscyphus fraternus (Peck) Dennis, Persoonia 3: 76, 1964,

NOTES. Helottum fraternum was originally described from
New York State as occurring on leaves, and was accepted by

ETS TEs acetone al ee Ses

|
|
|

White (1942). Dennis (1964) transferred the species to Hymen-
oseyphus; Arendholz (1979) did not treat the species. Dumont
(1981) redescribed and illustrated the species. He found that
the species formed a well-defined substratal stroma, produced
an ectal excipulum with hyphae embedded in a gelatinous matrix
and transferred the species to Poeculum (Sclerotiniaceae). For

peeuliedescraption, allustrations and discussion, see Dumont
(1981).

9. Helottum tmmutabtle Fuckel, Jahrb. Nassauischen Vereins
Na CUT Kees 52 Oe OO Om Gils FIGS.5.

= Hymenoscyphus tmmutabtlts (Fuckel) Dennis, Persoonia 3: 76, 1964.

Apothecial morphology — Apothecia scattered, arising from leaf blades,
short stipitate, ca 1.0-1.5mm in diam, to ca 0,5-0.75mm high, when fresh
flat to slightly convex, drying same, rehydrating flat to slightly cupu-
late, Hymenium when fresh white, drying pale yellow, yellow-brown, or
reddish orange, rehydrating slightly lighter and pallid; margin concolor-
ous with hymenium; receptacle when fresh white, drying pale yellow-orange,
rehydrating lighter and concolorous in the lower portion with the upper
portion of the stipe; stipe in the upper portion similar to receptacle,
coloration difficult to observe owing to small size,

Apothecial anatomy -—- Asci (70-)80-100(-105)x(6-)8-9um, produced from
small croziers, clavate to clavate-cylindric, gradually tapering toward
the base and there becoming expanded to form a small foot; wall ca lum
wide; pore J+. Ascospores (7-)8-10(-12)x2.5-3.5(-4.5)um, generally bise-
riate, but occasionally uniseriate, hyaline, smooth, obovoid to trapezoid-
al, anterior end round or rarely pointed, posterior generally slightly
pointed, in outline more or less equilateral, if inequilateral only
slightly so and not flattened on one side, walls slightly thickened, egut-
tulate or less commonly with polar guttulate areas (which are generally
obscured and difficult to detect) or in youth biguttulate with guttules
disappearing with age. Paraphyses equal to or slightly exceeding the
asci, internally hyaline, branching toward the base of the asci, septate,
filiform, becoming slightly expanded at the apex and there 2-3ym wide,
walls thin, smooth and hyaline. Subhymenium not well differentiated from
the medullary excipulum, consisting of parallel, vertically oriented hy-
phae grading into medullary excipulum. Medullary excipulum well devel-
oped, non-refractive, hyaline to pigmented light brown, consisting of sep-
tate, branched, parallel or loosely to tightly interwoven hyphae 2-4ym
wide, the walls thin, non-refractive, hyaline or rarely pigmented light
brown, and smooth. Ectal excipulum: inner ectal excipulum poorly defined
and differentiated from medullary excipulum, entire layer non-refractive,
hyaline to pigmented light brown, consisting of tightly compact, more or
less parallel, hyaline hyphae 2-3um wide, the walls thin, non-refractive,
hyaline and smooth. Outer ectal excipulum non-refractive, entire layer
hyaline to subhyaline, variable to ca 35um broad toward the margin and the
same toward the stipe, at the intersection of the stipe and the receptacle
to the outside composed of thin-walled, globose cells comprising a well
developed textura globulosa, to the inside the hyphae parallel, above the
juncture the isodiametric cells giving way to hyphae originating in the
stipe and extending almost perpendicularly or at very high angles to the
surface, progressing toward the margin the hyphae extending at lower and
lower angles to the surface until at the margin the hyphae parallel to the
surface of the apothecium, the individual cells with walls thin to slight-

ly thickened, hyaline or rarely pigmented light brown and smooth. Outer
covering layer as a distinct layer absent, but with an area to the outside
formed of the apical cells of the hyphae extending at low to high angles
to the surface and continuing to the surface, and then becoming somewhat
expanded and frequently large and clavate, these cells hyaline to slight-
ly yellow-brown with walls thin to slightly thickened and frequently
roughened. Margin constructed similarly to the upper portion of the
flank, with the individual cells smaller, the apical cells generally un-
modified. Hairs absent on the receptacle and margin. Stipe in the upper
portion constructed similarly to the lower portion of the receptacle, be-
low to the outside the cells globose with parallel narrow hyphae to the
inside; hairs absent.

Habitat: Leaves of Populus tremula, P. ntgra, Quercus pedunculata,
Quercus sp., Ulmus sp., Robinia pseudo-acacta.

Etymology of the specific epithet: relevance to the fungus uncertain.

Holotype: Germany, Boss Pr. Ebenbach, ad Popult tremulae folia putrida,
Fuckel (Fuckel, Fungi rhenani no. 2388, NY probably distributed in Herbier
Barbey-Boissier no. 1215.).

Tilustrations: Dennis, Mycol. Pap.62:°93, fig.) 85.) 1950 me Nn ecum rans
lowia 1: 143, fig. 3. 1943. Arendholz, Morphologisch-taxonomische Unter-
suchengen an blattbewohenden Ascomyceten aus der Ordnung der Helotiales,
Die bin etl OS me 2 ost 7 OR

nopres. SAccordins to: White *(1943) “andwArendholza(lo7o 1m
Helottum tmmutabtle is a widely distributed species in Europe
Dennis (1964)* transferred the species to Hymenoscyphus, a de-
cision with which I agree, and it appears most» closely relatea
tO Helottum epiphytlum, since? both havevsimilares truccuiesor
the sterile tissue of the apothecium; the outermost layers of
the stipe and base of the receptacle are composed of globose
cells, and the receptacle is composed of hyphae originating at
the stipe and at the base extending almost perpendicularly to
the surface. Further, from the stipe to the margin the hyphae
extend at progressively lower angles until toward the margin
the hyphae are parallel to the surface.

I have studied, few collections of #.; tmmutabctleswbuts based
on the shape of jthe ascospores, it appears distinets comer.
eptphyllum. There is overlap in the measurements, dH. epitphyl-
Zum 12-24x3-5ym (combined measurements according to Dennis,
1956, White, 1943 and Arendholz, 1979) and in #. tmmutabile
they are said to be 10-13x4-5um.. It%would) appear that asco-
sporen size is of little value in separating these two spe-
cies. I have attempted to locate the type specimen of Peziza
eptphylla in the Persoon herbarium at L, but no apothecia re-
main on the leaf, and all that is present in the type are sev-
eral apparently unrelated sclerotia, probably non-Sclerotinia-
ceae. Thus, a final decision on the correct placement of #.

FIG. 5. Hymenoseyphus immtabtilis, Fuckel, Fungi rhenani 2388 ex NY, free-
hand drawings, A, Median longitudinal section through an apothecium show-
ing lower portion of receptacle and margin, x 1,000, B, An ascus with 8
ascospores, C. 6 ascospores drawn after discharge from ascus, x 1,000,

D. Habit sketch of an apothecium on the substrate, x ca 50,

eptphyllum and its relationship with H. tmmutabile can be made |
only after a neotype specimen is designated for Peztza ept-
Divl layed clr es eam eno Canow prepared to do.

10. Helottum lindert White, Farlowia 1: 154. 1943.

vores. White described Helotium lindert from collections
from. the U.S.A. Dennis. (1964) ‘didenot *tream™ themspecies,
while Arendholz (1979) placed H. linderi into synonymy with
Hymenoscyphus caudatus, a decision with which I concur.

Holotype: U.S.A., Tenn., Chimney Trail, Great Smokey Mts. Nat. Park,
3200, ft. on fallen leaves, #18 Aug 1959.5. D. 7H slinders (En).

11. Helottum midlandense White, Farlowia 1: 605. 1944.
FIG...6%

Stroma — Substratal, visible on the fruits only as blackened areas of
the substrate; in section the blackened areas composed of an irregularly
formed rind composed of irregular to epidermoid cells in face view, also
visible in cross section of the base of the stipe of the apothecium; not
known in culture,

Apothecial morphology — Apothecia solitary, gregarious or occasionally
arising in clusters of 3-7, stipitate, unknown in fresh condition, when .
dry ca 0.5mm in diam and ca 0.5mm high, disc drying flat to slightly cupu-
late, rehydrating flat and expanded, Hymenium drying shades of yellow to
flesh-colored, rehydrating glassy-translucent; margin drying lighter than
hymenium, rehydrating concolorous with hymenium; receptacle drying concol-
orous with the margin, lighter than hymenium, rehydrating glassy-translu-
cent and concolorous with hymenium; stipe concolorous with receptacle in
dry and rehydrated conditions, cylindrical, short, generally less than 0.5)
mm high and wide, occasionally subpapillate in appearance.

Apothecial anatomy — Asci 8-spored, (40-)52-58(-65)x(5-)6-7um, pro-
duced from small, replicating croziers, cylindric to slightly clavate,
gradually tapering toward the base and there becoming slightly expanded to
form a small foot, wall to ca lum thick, slightly enlarged at the apex and
there to ca 2um thick; pore apparently J- in Melzer's Reagent. Ascospores
(S-)6-8(-9)x2-3um, obliquely uniseriate, irregularly biseriate or biseri- —
ate throughout, hyaline, smooth (a few devoid of pigment possibly lightly
punctate), aseptate, obovoid to obpyriform, ends rounded or rarely slight- ~
ly pointed, in outline generally equilateral, anterior end obviously
broader than posterior end, occasionally slightly indented below median
point of spores; guttules generally absent or rarely with one tiny (ca lum
or less), transversely positioned at the narrowest portion of the spore
and appearing as a septum. Paraphyses equal to or slightly exceeding the
asci, internally hyaline, branching toward the base of the asci, septate,
cylindrical, generally not becoming expanded at the apex and there 2-3ym
wide, walls thin, smooth, and hyaline. Subhymenium well developed, well

BIG, 6. Helottum mtdlandense, Stevens 59 ex NY, freehand drawings, 1,000,
A. Median longitudinal section of an apothecium through margin, B, Median ~
longitudinal section of an apothecium through the juncture of receptacle

and stipe. C. An ascus with 8 ascospores and a paraphysis. D. 9 asco-
spores drawn after discharge from ascus.

differentiated from the medullary excipulum below, hyaline, to ca 50um 4
broad in the center, narrower toward the margin, consisting of parallel to,
slightly interwoven, vertically interwoven hyphae, the individual hyphae
hyaline, 1.5-3um wide, the walls thin, hyaline and smooth. Medullary ex-
cipulum poorly developed, obconical, non-refractive, hyaline, consisting
of septate, branched, loosely to tightly interwoven (to parallel in the
flank) hyphae 2-3(-4)um wide, the walls thin, hyaline and smooth. Ectal
excipulum: inner ectal excipulum well to poorly defined, well-differenti- |
ated from the outer ectal excipulum and grading into the medullary excipu- |
lum, entire layer non-refractive, hyaline, 15-20um broad toward the margin |
and approximately the same toward the stipe, consisting of parallel to
slightly interwoven, hyaline hyphae 2-3um wide, the walls thin, non-re-
fractive, hyaline and smooth. Outer ectal excipulum non-gelatinized, non-|
refractive, entire layer hyaline, toward the stipe ca 50yum broad, consist-|
ing of large, globose, angular to irregular cells forming a well defined |
textura globulosa to angularis, the individual cells (5-)8-12(-20) um wide, |
the walls thin to slightly thickened, hyaline and smooth. Outer covering
layer and hairs absent; progressing towards the margin the outer ectal ex- |
cipulum becoming narrower, and grading from a textura angularis and globu- |
losa to textura prismatica, the individual cells smaller and forming more |
and more regular rectangularly shaped cells until at the margin the layer
composed of a well defined textura prismatica, lacking an outer covering
layer, Margin simple, the brick-shaped cells if continuing into margin
only a short distance and the remainder simple and composed of the narrow |
cells of the inner ectal excipulum and the adjoining paraphyses, hairs |
also absent in the margin. Stipe constructed similarly to the lower por- |
tion of the receptacle, to the outside a zone to ca 70Oym broad composed of |
globose to angular cells and grading into narrow, parallel hyphae in the
interior of the stipe; the individual ‘cellsiwith wallssthingtossligngly
thickened, hyaline and smooth or rarely roughened; at the base rind cells
visible; hairs absent.

Habitat: Old pods of Gledttsta trtacanthos L. and reported by White
(1944) to occur also on petioles and midveins of Quercus sp.

Etymology of the specific epithet: refers to the part of the country
where the fungus was originally collected.

Holotype (not examined): Iowa, Homestead, pods of Gledttsta trtacan-
thos, 26 Sept 1931, G. W. Martin 5186 (FH).

Illustrations: White, Farlowia 1: 607, "figs. 10-13. 1944"

Specimen examined: Kansas, Lawrence, on pods (? of Gledttsta sp.),
1890, W. C. Stevens 59 (paratype ex NY).

NOTES. When White (1944) described Helotium midlandense,
he noted that the stipe to the outside was composed of globose |
to angular cells, and he illustrated the ectal excipulum as
formed of brick-shaped cells in the lower portion approaching
the stipe. I have examined a paratype collection and concludem
that the majority of the ectal excipulum is composed of a well
defined textura globulosa to angularis, but that toward the
margin the tissue becomes more organized into a well defined
textura prismatica, and not toward the stipe as illustrated by
White. Further, I have found a well developed rind on the
substrate in association with the apothecia and in section
have observed rind cells at the base of the stipe. I have
also noted that the spores are relatively small for the Scle-

1s)

otiniaceae and are oboyoid to obpyriform, Since the species
Peoduces arsubstrata le stroma. 1t should be referred to the
pelenoviniaceae, —iBecause (i)) the ectal; excipulum is composed
of a textura globulosa, (ii) small pyriform ascospores are
present, and (111) it occurs on fruits, the species is prob-
Bplybest referred Co Ciboria. At present, J] am- uncertain of
species concepts in Ctborta and of the separation between Ci-
borta and Moellerodtseus; I will not make a new combination

in Ctborta, but will defer action until the species are better
known.

I have observed in several apothecia a rather high propor-
tion of misshapen and malformed ascospores, some of which pro-
duced an apparent septum. I cannot satisfactorily explain the
presence of the irregularly formed ascospores, nor the occar
sional production of a single septum.

My observations are then different from White's, who did
not» report the presence of a stroma and referred it to the
Leotiaceae (Helottum). Neither Dennis (1964) nor Arendholz
(1979) treat the species.

ice Herortum pnytlogenon Rehm, Hedwigia 24: 14. 13885.

= Hymenoscyphus phyllogenon (Rehm) Kuntze, Revis. gen, Pl, 3(3): 485,
TSOse

NOTES. Helottum phyllogenon was originally described from
Hungary on leaves of "poplar." Kuntze, (1898) transferred the
species to Hymenosecyphus, and the placement there was accepted
Dy Dennis (1964) and by Arendholz (1979). I find the type
collection to be indistinguishable from #. caudatus, following
Poewconcept Of Dumont Gg. Carpenter (1981). They gave the asco-
PoOore Measurements O12. caudatus to be (14-)16-23(-26)x4-5
(-6)ym, whereas in the original description of H, phyllogenon
the :ascospore measurements were 12-15x3.5um. White (1943)
gave them as 11-14x3.8-5um, Dennis (1956) 14-16x4-6um, Arend-
Zc, oye oxo. OAM. tL tind =the apothecial “structure
of the type of H. phyllogenon indistinguishable from H. cauda-
tus, and conclude that they represent the same species, with
the ascospore measurements of the type of H. phyllogenon fall-
ing at the lower limits of #. caudatus and perhaps extending
the limits somewhat lower than previously reported by Dumont §
Barpenter (19381 )*,

Holotype: Hungary: bei Ungerisch-Altenburg (Ungarn), an faulen Pallel-
Blattern, Oct 1883, Linhart (ex FH, Rehm, Ascomyceten 768).

13. Helotium phyllophilum (Desmaziéres) Fries, Summa veg.
Scandi. 2500. 1849.

Peziza phyllophtla Desmaziéres, Ann., Sci. Nat. Bot., Sér. 2, 17:
98. 1842; PZ. Crypt (du Nord) France éd. 1. no. 1159. 1842; éd.
eiser- leno. 659. 91842.

= Hymenoscyphus phyllophtlus (Desmaziéres) Kuntze, Revis. gen. Pl.

3(3): 485. 1898.

NOTES. ‘Peziza phyllophila was originally described as oc-
curring on leaves of Acer and Fagus from France and was trans-

ferred to Helottum by Fries and to Hymenoscyphus by Kuntze
(1898). Its placement in Hymenoscyphus was accepted by both
Dennis (1964) and Arendholz (1979). I find the sterile tissue#
of the apothecium of the lectotype specimen of the species to]
be indistinguishable from that of H. caudatus. The apothecia
of H. phyllophilus were originally thought to be smaller than
those of H. caudatus, but as Dumont §& Carpenter have pointed
out, the apothecia of tropical collections are frequently less@
than lmm in diameter and those of the type of H. phyllophilus |
would then fall within the range of H. caudatus.

In the original description the measurements of the asco-
spores were not given, but they were reported to be 1-septate.}
Dennis (1956) reported the ascospores to be 14-16x3.5-4um,
White (1943) gave them as 11.5-15x3.2-4ym, and Rehm (1893, in
1887-1896) had them 10-15x3-3.5ym. Rehm further placed Helo-
tium phyllophtlum and H. phyllogenon into synonymy. The only
difference which I consider noteworthy between Hymenoscyphus
caudatus and H..phyllophilus is.that in the latter aliotache
ascospores in the type collection are l-septate, whereas, as
pointed out by Dumont §& Carpenter (1981), in H. caudatus they
are.only rarely l1-septate. However, I have noted in other
clearly marked species, such as H. rufocorneus (see Dumont,
1981) where for some unexplained reason all of the ascospores
in a single collection become 1l-septate. I do not consider |
this then to represent the single criterion upon which to sep#
arate two species, H. caudatustand) A. phyblophilus. s hatnet as
prefer to consider them as probable synonyms.

There 1S some question as to where, the name fe2z7zaquony lo
phittla" was actually first published wand as tos whagetnescype |
specimen actually is. Desmazi@éres (1842) presented a formal
description of the species and a description also appeared on
the labels of two specimens cited by Desmaziéres; "Pl. Crypt. @
éd. J, no. 1159; éd. 2, no. 659.'"" As nearly as (can, be (detcr am
mined (Sayre, 1969), fascicles containing these two specimens
were issued simultaneously. I have been unable to determine
if the article appeared before the exsiccati specimens.

In this regard, White (1943) selected no. 1159 as the type,
but indicated that he examined two specimens, one in the bound)
set of the exsiccata at FH and the other from the Curtis col-@
lections. I select the portion deposited in the bound exsic- @
cata as the lectotype specimen for Péztiza phytlopnitla.

Lectotype specimen: Plantes Cryptogames de France, éd. 1, no. 1159 (ex
FH, bound set).

14. Helottum translucens White, Farlowia 1; 149. 1943.

FIG. 7. Hymenoscyphus transtlucens, holotype ex FH, freehand drawings,

x 1,000. A. Median longitudinal section of an apothecium through margin.
B. Median longitudinal section of an apothecium through margin. C. An
ascus with 8 ascospores. D. A branching paraphysis. EE. 8 ascospores
drawn after discharge from ascus.

= Hymenoscyphus translucens (White) Arendholz, Morph.-tax. Untersuch.
blattbe. Ascomyceten Helotiales p. 78. 1979.

Apothecial morphology — Apothecia tiny, solitary to occasionally gre-
garious, arising from leaf blades, especially veins of various sizes, with
a minute stipe, to ca 0.5mm in diam and to ca 0.4mm high, when fresh disc
flat, drying and rehydrating same. Hymenium pallid to hyaline when fresh, |
drying off-white to pale ochraceous, rehydrating lighter, pale flesh-col-
ored to off-white; margin concolorous with hymenium when fresh, dry and
rehydrated; receptacle generally concolorous with the hymenium when fresh, |
dry and rehydrated; stipe, difficult to observe coloration owing to minute |
Size. |

Apothecial anatomy — Asci 8-spored, 80-95x9-llum, probably produced
from tiny croziers, long cylindric-clavate, gradually tapering toward the
base and there not expanded to form a small foot, wall ca lum wide,
slightly enlarged at the rounded to subtruncate apex and there 2(-3)um
thick; pore J+. Ascospores (10-)11-14(-15)x(3-)4(-5)um, uniseriate to bi-|
seriate throughout, hyaline, smooth, aseptate or very rarely l-septate,
trapezoidal, obovoid, ends rounded or less commonly slightly pointed, in |
outline generally equilateral, if inequilateral only slightly so and then
not flattened on one side, anterior end broader than posterior end, fre-
quently appearing eguttulate or with two polar guttulate areas generally |
obscure but granular and oily in composition, walls slightly thickened and
occasionally to ca lym wide. Paraphyses equal to or slightly exceeding |

the asci, internally hyaline, branching at the base of the asci and toward™

the middle of the asci, filiform, becoming slightly expanded at the apex |
and there (2-)3-4ym wide, walls thin, smooth and hyaline. Subhymenium not”
well differentiated from the medullary excipulum, but with a tendency of ||
the hyphae at the base of the asci to be slightly more compact and narrow-_
er than the hyphae in the center of the receptacle. Medullary excipulum
poorly developed, obconical, non-refractive, hyaline, consisting of .sep-
tate, branched, more or less parallel to slightly interwoven hyphae 2-4
(-5)um wide, the walls thin, non-refractive, hyaline and smooth. Ectal
excipulum: inner ectal excipulum poorly defined and differentiated from
medullary excipulum. Outer ectal excipulum non-refractive, non-gelati-
nized, entire layer hyaline to subhyaline, ca 15-20ym wide toward the mar-_
gin and to ca 20-25ym toward the stipe, consisting predominantly of tex- |
tura prismatica with the individual hyphae extending parallel to or at low
to high angles to the surface of the apothecium and without apically free
hyphal tips; the individual cells frequently collapsing, becoming disori-
ented owing to drying process and tissue then appearing to be composed of
narrow hyphae; individual cells toward the margin 5-12(-15)x3-5ym and 12-
22x3-7ym toward the stipe, the walls thin to slightly thickened and ca 1
um broad, hyaline to light brown and smooth. Outer covering layer pres-
ent, but difficult to detect, 1-2 hyphal layers and 1-3um broad, the in-
dividual hyphae extending parallel to the surface of the apothecium, over- |
lapping, terminating before the margin, the individual cells subhyaline,
the walls non-refractive, thin, pigmented light brown to intense brown,
smooth to frequently slightly roughened. Hairs absent. Margin poorly de-
veloped, narrow above, broader below, entire zone hyaline to light brown,
constructed similarly to the flank below, the individual cells smaller.
Stipe in the upper portion constructed similarly to the lower portion of
the flanks, not becoming noticeably interwoven in the center, toward the
base the individual hyphae becoming disrupted, and frequently losing hy-

phal orientation, but generally oriented at a high angle to the surface.
Hairs absent.

Habitat: Leaves of Acer sp., Quercus sp. and Fagus stlvattca (fide
Arendholz, 1979).

Etymology of the specific epithet: refers to the color of the apotheci-
um in its fresh condition.

Holotype: U.S.A., Mass., swamp at east side of Great Blue Hill, near
Milton, on decaying leaves of Acer sp., 11 Nov 1941, D. H. Linder, E. V.
seeler G W. L. White s.n. (FH).

Paratype: U.S.A., Mass., Stony Brook Reservation, West Tuxbury, fallen
leaves of Quercus sp., 11 Nov 1941, W. L. White s.n. (FH).

Pilustracions: White, Farlowia 1:31595,, £19. 6..1943- Arendholzs Mor-
phologisch-taxonomische Untersuchungen an blattbewohenden Ascomyceten aus
MoreHelotrales plomi0, L19S: 5, eo.unl 9/9.

NOCD OMe NentS deSCEINDCION Ole HeLOL TUM Lrane ucenss White
(1943) concluded that this new species was closely related to
Helottum caudatum, and I am in agreement with this. dHelottum
Pauaarum 1S) dadistinguashnedy trom Hi translucens in®that the apo-
mneCiavOt A. evrransilucens, are smaller than those of f° *caudatum
and the ascospores in H#:-caudatum are predominantly ‘inéequilat-
erateancetrequenctly Nnooked apically sn) contrase the spores of
Pomcrane pucenceare trapezoldalwin shape: andsgenerally equilat-
eral. s lhe shape of the ascospores in”. translucens 1S sug=
PoctiViewOt ChOSCmOLa-m7Tmnicap1 cow mine Structure: OL the apo-
thecia in these two species is fundamentally different. See
discussion under H. tmmutabtle for a comparison of apothecial
structure.

PhemspecresmWwasenO tac reatedebysDenn1s. (1956,.1904).,, butt
accept the placement of #. translucens in Hymenoscyphus by
mrendholz (¢1979)..

Acknowledgements

anesauthorawisnesto express is gratitude, to the directors of those
institutions who have provided the loan of material for the present study,
injparticular, Dr. D. H. Pfister, of FH has been most helpful in the loan
of specimens and assistance in working out bibliographic citations. Fi-
nancial support for the present study has been provided by the National
Science Foundation grant, GB. 28593. The line drawings have been prepared
by Bobbi Angell.

LITERATURE CITED

Arendholz, W.-R. 1979. Morphologisch-taxonomische Untersuchungen an
blattbewohenden Ascomyceten aus der Ordnung der Helotiales. A disser-
tation presented to the faculty of Hamburg University. 1-115 pp., pl.
1-16.

Carpenter, S. E. 1981. Monograph of Croctecreas (Ascomycetes, Helotiales,
Leotiaceae). Mem. New York Bot. Gard. 32: 1- in=press.

& K. P. Dumont. 1978. Los Hongos de Colombia-IV. Bisporella trisep-
tata and its allies in Colombia. Caldasia 12(58): 3359-348,

Dennis, R. W. G. 1956. A revision of the British Helotiaceae in the her-
barium of the Royal Botanic Gardens, Kew, with notes on related Europe-
anpspectes, e Mycol. ap., 02251-2116, apl 1.

1964. Remarks on the genus AHymenoscyphus S. F. Gray, with observa-

tions on sundry species referred by Saccardo and others to the general

Helotium, Peztzella or Phtalea. Persoonia 3: 29-80.

Desmaziéres, J. B. H. J. 1842. Cryptogames nouvelles. (Ann. Sci 7sNat.
Botewsoen 2 salir la Liss

Dumont, K. P. .1971. Sclerotiniaceae II. Lambertella. Mem. New York Bot.
Gard’: 22(4) te-1=7 3"

1976. Sclerotiniaceae XI. On Moellerodiscus. Mycologia 68: 233-
PLONE
1980. Sclerotiniaceae XVI. On Helotium rufo-corneum and Helottum
fraternum. Mycotaxon 12(1): 255-277.
1981. Leotiaceae II. A preliminary survey of the neotropical

species referred to Helottum and Hymenoscyphus. Mycotaxon 12(2): 313-
SAG

& S. E. Carpenter. 1981. Leotiaceae IV. Los Hongos de Colombia-VII.
_ Hymenoscyphus caudatus and related speci¢s from Colombia and adjacent
regions. Caldasia (in press).

Haines, J. H. 1980. Studies in the Hyaloscyphaceae 1: Some species of
Dasyscyphus on. tropical ferns. Mycotaxong ll 31389-21673

Kuntze; 0; 18984) sReviss gen. Pl 445(3)? Gl-576-

Peck; C.,H#) 1879. Reportof the botanist.) Annual) Rep. New, Yorkeotate
Musii5517 19-60.

Rehm, H. 1887-1897. Die Pilze. Ascomyceten: Hysteriaceen und Discomy-
ceten. In Rabenhorst's Kryptogamen-Flora von Deutchland, Oesterreich
Undederecchwre zee cenUllawico) ela 75.

Sayre, G. 1969. Cryptogamae Exsiccatae - An annotated Bibliography of
Published Exsiccatae of Algae, Lichens, Hepaticae, and Musci. Mem. New
LOTKBOtUs Gard 1619 (0) cme Lae

White, W. L. 1942. Studies in the genus Helottum, I. A review of the
species described by Peck: Mycologia 34: 154-1797

1943. Studies in the genus Helottum, III. History and diagnosis of
certain European and North American foliicolous species. Farlowia 1:
135-170.

1944. Studies in the genus Helottum, IV. Some miscellaneous spe-
cies meebariowiarl: 8599-6172

MYCOTAXON

Vode ALIS No eal epp. S024 04 April-June 1981

LICHENES EXSICCATI
DISTRIBUTED BY THE UNIVERSITY OF COLORADO
MUSEUM, BOULDER
FASCICLES 1-15, NOS. 1-600, 1961-1979

WILLIAM A. WEBER

Univ. of Colorado Museum,
Campus Box 218, Boulder, 0 80309

SUMMARY

Revision of identifications and nomenclature
for the first 600 numbers of LICH. EXSICC. OOLO is
provided, and the following new combinations are
proposed: Cladina galapagosensis, C. polia, Hetero-
dermia barbifera, H. circinalis, H. stellata and H.
verrucifera. Validation is provided for the publi-
cation of three new species distributed in the ex-
siccati: Lecanora pseudopinguis, L. texana and Psora
cerebriformis.

Lichenes Exsiccati, distributed by the University of Colorado Museum
(standard abbreviation LICH. EXS. COLO) began in 1961 with the distribu-
tion of Fascicle 1 (1-40). The latest fascicle was Fascicle 15 (56]-
600), distributed in 1979. Some of the early fascicles were distributed
in 100 sets, but the current ones are reduced to 60 sets. The exsiccati
are offered in exchange to most of the active lichen herbaria of the
world. A brief notice was published in Taxon 13:31. 1964, wherein the
institutions receiving the complete series were listed. The present pa-
per provides an alphabetical arrangement of the numbers thus far distri-
buted, with updated nomenclature and corrections of identifications up to
the present moment. Additional corrections will be welcomed.

In the following list, the first name given is that under which the
number was distributed. When that name is enclosed by brackets, this in-
dicates that the name has been changed, amd the current name is preceded
by an (=) sign. If the identification has been changed, no (=) sign pre-
cedes the second name.

TAXON

Acarospora badiofusca (Nyl.) Th. Fr.

[A. flava (Bell.) Trev.] =A. chlorophana (Wg.) Ach.

A. nitida H. Magn.

A. schleicheri (Ach. ) Mass.

[Alectoria chalybeiformis (L.) S. Gray] =Bryoria
chalybeiformis (L.) Brodo & Hawksw.

[A. fremontii Tuck.] Bryoria tortuosa (Merr.) Brodo
& Hawksw.

[A. glabra Mot.] Bryoria lanestris (L.) Brodo &
Hawksw.

A. lata Tayl.

A. nigricans (Ach.) Nyl.

A. ochroleuca (Hoffm.) Mass.

A. poeltii Bystrek

[A. pubescens (L.) R. H. Howe] =Pseudephebe pubes-
cens (L.) Choisy

A. sarmentosa (Ach.) Ach.

A. smithii DuRietz

[A. tenuis E. Dahl] Bryoria chalybeiformis (L.) Brodo
& Hawksw.

[Anaptychia barbifera (Nyl.) Trevis. ] =Heterodermia
barbifera comb. nov.

[A. circinalis (A. Zahlbr.) W. A. Web.] =Heterodermia

circinalis comb. nov.
[A. diademata (Tayl.) Kurok.] =Heterodermia diadem-

ata (Tayl.) Awasthi

A. erinacea (Ach.) Trevis.

[A. leucomela (L.) Mass. f. verrucifera Kurok. ]
=Heterodermia verrucifera comb. nov.

[A. multiciliata Kurok.] =Heterodermia multiciliata
Follm. & Redon

[Anaptychia obscurata (Nyl.) Vain.] =Heterodermia
obscurata (Nyl.) Trevis.

[A. stellata (Vain.) Kurok.] =Heterodermia stellata
(Vain.) W. A. Web., comb. nov.

Anthracothecium ochraceoflavum (Nyl.) Muell.-Arg.

Anzia angustata (Pers.) Muell.—Arg.

A. gregoriana Muell.—Arg.

A. hypoleuca Muell.-Arg.

A. wilsonii Raes.

Arthonia glebosa Tuck.

Norway

California

Wyoming

California

Tasmania
Canada
N. Guinea

Canada
Norway

California

N. Guinea
Colorado
N. Guinea
Galapagos
Arizona
Mexico
Mexico
Galapagos
Chile
Australia
Galapagos
Galapagos
Australia
N. Guinea
N. Guinea

Australia
Colorado

p35

508
145
281
377
378
311

A. impolita (Ehrh.) Borr. ex Hook. & Sowerby

A. platyspilea Nyl.

A. rubella (Fee) Nyl.

Arthopyrenia halodytes (Nyl.) Arn.

[Arthothelium spilomatoides (Nyl.) A. Zahlbr.]
A. galapagoense Huneck & Follmann TYPE OOLL.

Aspicilia alpina (Sommerf.) Arn.

A. desertorum (Kremp.) Mereschk.

[A. hispida Mereschk.] A. desertorum (Kremp.)
Mereschk.

[A. mutabilis (Ach.) Koerb.] =Pachyospora mutabilis

(Ach. ) Mass.
A. quartzitica W. A. Web. TYPE COLL.
A. transbaicalica Oxner
A. sp. indet. (fruticose modification)
Bacidia albescens (Arn.) Zwackh.
B. herrei A. Zahlbr.
B. millegrana (Tayl.) A. Zahlbr.
Baeomyces absolutus Hepp in Zollinger

B. trachypus Nyl.

B. weberi J. W. Thomson TYPE OOLL.
Belonia americana Fink ex Hedrick

Bryoria tortuosa (Merr.) Brodo & D. Hawksw.
Buellia curtisii (Tuck.) W. A. Web.

B. flavoareolata (Nyl.) Muell.-Arg.

B. galapagona W. A. Web. TYPE COLL.

B. glaziouana (Kremp.) Muell.-Arg.

B. novomexicana B. de Lesd.

B. oidalea (Tuck.) Tuck.
B. punctata (Hoffm.) Mass.
[B. spuria (Schaer.) Anzi]
undescribed taxon.

. cf. taltalensis Dodge

. triphragmioides Anzi
sp. indet.

. Sp. indet.

. zahlbruckneri Steiner
Calicium abietinum Pers.
[C. hemisphaericum Howard] =C. adequatulum Nyl.
Caloplaca amabilis A. Zahlbr.

C. aurantiaca (Lightf.) Th. Fr.

Incorrect, possibly an

| Gd) Oo) |

[C. bracteata (Hoffm.) Mass.] Fulgensia desertorum

(Tomin) Poelt, cited specimen!
[C. chrysophthalma Degel.] Determination doubt ful.
C. cinnanomea (Th. Fr.) Oliv.

California
California

USSR
California
California
Oregon
Galapagos
N. Guinea
N. Guinea
N. Guinea
N. Guinea
Texas
Canada
Louisiana
Chile
Galapagos
Galapagos
Arizona
California
Colorado

Mexico
Chile
Colorado
Peru
Peru
Arizona
Idaho
Montana
Arizona
Arizona
Colorado

N. Mexico
Colorado

C. cladodes (Tuck.) A. Zahlbr. in Engler

C. epithallina Lynge

C. fernandeziana (A. Zahlbr.) Follm. & Redon

C. lamprocheila (DC.) Flagey

C. microphyllina (Tuck. ) Hasse

C. modesta (A. Zahlbr.) Fink

C. pollinii (Mass.) Jatta

C. stanfordensis H. Magn.

C. subnitida (Malme) A. Zahlbr.

C. trachyphylla (Tuck.) A. Zahlbr.

[Caloplacopsis submexicana (B. de Lesd.) B. de Lesd. ]
=Candelina submexicana (B. de Lesd.) Poelt

Candelariella deflexa (Nyl.) A. Zahlbr.

C. rosulans (Muell.—Arg.) A. Zahlbr.

C. spraguei (Tuck.) A. Zahlbr.

Catillaria griffithii (Sm.) Malme

C. sp. indet.

Cavernularia lophyrea (Ach.) Degel.

Cetraria canadensis (Raes.) Raes.

C. cucullata (Bell.) Ach.

C. delisei (Bory ex Schaer.) Th. Fr.

C. ericetorum Opiz

C. fendleri (Nyl.) Tuck.

[C. fendleri f. coralligera W. A. Web. TYPE OOLL.]
=C. coralligera (W. A. Web.) Hale

[c. glauca (L.) Ach.] =Platismatia glauca (L.)
W. Culb. & C. Culb.

C. idahoensis Essl.

C. islandica (L.) Ach.

C. pinastri (Scop.) S. Gray

[C. richardsonii Hook. in Richards.] =Masonhalea rich-

ardsonii (Hook. ip Richards.) Kaernefelt

C. scutata (Wulf.) Poetsch

C. tilesii Ach.

C. sp. indet.

Cetrelia chicitae (W. Culb.) W. Culb. & C. Culb.

Chaenotheca brunneola (Ach.) Muell.~Arg.

Chiodecton effusum Fee

C. farinaceum Fee

C. sp. indet.

Chondropsis semiviridis (F, Muell. ex Nyl.) Nyl. ex
Cromb. i ae

Colorado
Colorado
Chile
Colorado
Arizona
Arizona
Minnesota
California
Texas
Colorado

Arizona
Colorado
Colorado
Colorado
Colorado
California
Chile
Canada
Idaho
Alaska
Norway
Colorado
Arizona

Arizona

Colorado
Idaho
Colorado
Australia
Colorado

Alaska
Tasmania
Alaska

N. Guinea
N. Guinea
Idaho
Galapagos
Galapagos
Chile
Chile

Australia
Australia

Cladia aggregata (Sw.) Nyl.

C. ferdinandii (Muell.-Arg.) R. Filson
C. retipora (Labill.) Nyl.

[C. retipora (Labill.) Nyl.] C. corallaizon R.
Filson

C. schizopora (Nyl.) Nyl.

C. sullivanii (Muell.-Arg.) W. Martin

[Cladonia alpestris (L.) Rabenh.] =Cladina stellaris
(Opiz) Pouzar & Vezda

[Cladonia cariosa (Ach.) Spreng. ]

C._ceratophylla (Sw.) Spreng.

C. chlorophaea (Flk.) Spreng.

C. coccifera (L.) Willd.

C, corymbescens Nyl.

C.e a Leighton

C. foliacea (Huds.) Willd.

[C. galapagosensis Ahti] =Cladina galapagosensis
comb. nov.

C. leporina Fr.

C. multiformis Merrill

C. robbinsii Evans

C. pertricosa Kremp.

C. cf. pityrea (Flk.) Fr.

[C. polia R. Sant.] =Cladina polia (R. Sant.) comb.
pOve’

C. scabriuscula (Del. in Duby) Nyl.

c. solitaria solitaria H. Magn.

TC. squamosa squamosa (Scop.) Hoffm.] C. dilleniana Flk. fide
J. W. Thomson ;

[C. subtenuis (des Abb.) Evans] =Cladina subtenuis
(des Abb.) Hale & W. Culb.

C.. sulphurina (Michx.) Fr.

C. verticillata (Hoffm.) Schaer.

C. vulcanica Zoll. in Hasskarl

[C. xanthoclada Muell.-Arg.] C. capitellata (Tayl.)
e Raby ee.

C. zopfii Vain.
Coelocaulon aculeatum (Schreb.) Gyel.

[C. australiense nom. nud. |
Kaernefelt TYPE COLL.
Coenogonium implexum Nyl.

Collema cristatum (L.) G. H. Web. in Wiggers

Cetraria australiense

Nepal

Australia
Galapagos
Australia
Australia

Australia
Australia
Australia

Tasmania

Alaska

Kansas
Galapagos
Galapagos
N. Guinea
N. Guinea
Colorado
Denmark

Galapagos
Alabama

Colorado —

N. Mexico
Australia
Australia

Galapagos
Australia
Chile

N. Guinea

Kansas
Montana
Australia
N. Guinea

Australia
Denmark
Colorado
Canada

Australia
Tasmania
Colorado

C. durietzii Degel.

C. leucocarpum Hook. f. & Tayl.

C. polycarpum Hoffm.

Conotrema urceolatum (Ach.) Tuck.

[Cora pavonia (Sw.) Fr.] =Dictyonema montanum (Sw. )
Marm. ex Follm.

[Cornicularia aculeata (Schreb.) Ach.] =Coelocaulon
aculeatum (Schreb.) Gyel.

[C. californica (Tuck.) DuRietz] =Coelocaulon cali-
fornicum (Tuck. ) Howe f.

C. nommerica (Gum. ) DuRietz

[Cypheliopsis bolanderi (Tuck.) Vain.] =Thelomma
mammosum (Hepp in Hartung) Mass. The collection
is mixed, containing also T. santessonii L, Tibell

Cyphelium inquinans (Sm.) Trev.

stocoleus ebeneus (Dillwyn) Thwaites
Dactylina madreporiformis (Ach. ) Tuck.

Darbishirella gracillima (Kremp.) A. Zahlbr.
Dendrographa leucophaea (Tuck. ) Darb.

D. minor Darb.

Dermatoc leptophyllum (Ach.) Vain.
D. miniatum Ta) Mann

D. plumbeum B. de Lesd.
D. reticulatum H. Magn.
D. rivulorun (Arn.) Dalla Torre & Sarnth.
D. tuckermanii (Rav. ex Mont.) A. Zahlbr.
D. vellereum Zschacke _

Dictyonema irpicinum Mont.

D. sericeum (Sw.) Berk.

Diploicea canescens (Dicks.) Mass.

Diploschistes ocellatus (Vill.) Norm.

[D. scruposus (Schreb.) Nomm.] D. cf. canadensis
Raes.

Dirina catalinariae Hasse

D. herrei A. Zahlbr.

D. limitata Nyl.

Dolichocarpus chilensis R. Sant.

Endocarpon pulvinatum Th. Fr.

E. pusillum Hedw.

Enterographa atacamensis Dodge

[Enterographa atacamensis Dodge] Roccellina olivacea

Follm.
Ephebe lanata (L.) Vain.
Evernia prunastri (L.) Ach.
EI ROBIE RICA EERE OE RR

Australia
Australia
Colorado

W. Virginia

Galapagos
Colorado

Oregon
Oregon

Mexico
Colorado
Colorado
Colorado
Colorado
Chile
Mexico
Mexico
Calif.
USSR
Arizona
Colorado
Oregon
Colorado
Arizona
USSR

N. Guinea

N. Guinea
California

Mexico

Colorado
Mexico
Galapagos
Chile
Chile
Colorado
Colorado
Colorado
Peru

Chile
Colorado

California

438

Everniastrum neocirrhatum (Hale & V. Wirth) Hale

E. vexans Hale

Glypholecia scabra (Pers.) Muell.-Arg.

Graphis caesiella Vain.

G. striatula (Ach.) Spreng.

Gyrostomum scyphuliferum (Ach. ) Nyl.

Haematonma babingtonii Mass.

H. subpuniceum (Muell.-Arg.) B. de Lesd.

Heppia lutosa (Ach.) Nyl.

Heterodea melleri (Hampe) Nyl.

Heterodermia leucomelos (L.) Poelt

[Hubbsia lumbricoides W. A. Web. TYPE COLL.]
=Reinkella californica Raes.

Hydrothyria venosa Russell

Hypogymnia billardieri (Mont.) R. Filson

H. krogii Ohlsson

H. lugubris (Pers.) Krog

H. mundata (Nyl.) Rassadina

H. oroarctica Krog

H. physodes (L.) Nyl.

Icmadophila ericetorum (L.) A. Zahlbr.

Ingaderia pulcherrima Darb.

Koerberia biformis Mass.

Lasallia pensylvanica (Hoffm.) Llano

L. pustulata (L.) Mereschk. ssp. papulosa (Ach. )

W. A. Web.

Lecanactis californica Tuck.

L. cf. myriadea (Fee) A. Zahlbr.

L. (subg. Bactrospora) sp. indet.

[Lecanora arizonica (Tuck.) W. A. Web.] =Omphalo-
dium arizonicum (Tuck. ) Tuck.

L. badia (Hoffm.) Ach.

[L. bolanderi Tuck.] lL. phryganitis Tuck.

[L. cancriformis (Hoffm.) Vain.] L. caesiorubella
Ach. ssp. merrillii Imsh. & Brodo

L. carpinea (L.) Vain.

L. chlarotera Nyl.

L. christoi W. A. Web. TYPE OOLL.

L. confusa Almborn

[L. conizaea (Ach.) Nyl.] Incorrect, but no alter-
native identification available.

[L. conizaeoides Nyl. ex Cromb.] L. confusa Almb.

L. coquimbensis A, Zahlbr.

L. frustulosa (Dicks. ) Ach.

L. garovaglii (Koerb.) A. Zahlbr.
ments

(Disregard com-

Mexico
Galapagos
USSR
Galapagos
Galapagos
Louisiana
New Zealand
Texas
Iowa
Australia
Mexico

Mexico
California
Australia
N. Carolina
Australia
Australia
Colorado
Colorado

Colorado
California
Chile
California

N. Mexico
Montana
California

California
California
Colorado
Colorado
Oregon

Galapagos
California

Chile
Colorado

Colorado

[L. cf. glabrata (Ach.) Malme] Probably incorrect
but no alternative name available.

[L. lentigera (G. Web.) Ach.] =Squamarina lentigera
(G. Web.) Poelt

([L. marginalis Hasse] =Rhizoplaca marginalis (Hasse)
W. A. Web.

[1.. melanophthalma Ram.] =Rhizoplaca melanophthalma
(Ran. ) Leuckert & Poelt

L. mellea W. A. Web. TYPE COLL.

[L. peltata (Ram.) Steud.] =Rhizoplaca peltata (Ram. )

Leuckert & Poelt

L. phryganitis Tuck.

L. pinguis Tuck.

L. polytropa (Ehrh.) Rabenh.

L. pringlei M. Lamb

L. pseudopinguis W. A. Web. sp. nov. TYPE COLL.

[L. rubina (Vill.) Ach.] =Rhizoplaca chrysoleuca
(Sm.) Choisy

L. saligna (Schrad.) A. Zahlbr.

L. texana W. A. Web., sp. nov. TYPE COLL.

[L. thomsonii H. ent =L. novomexicana B. de Lesd.
ex H. Magn.

L., stirps varia (Ehrh.) Ach.

Lecidea aspidula Kremp.

[L. atrobrunnea (Ram. ): Schaer. ]
Arn.

L. berengeriana (Mass.) Th. Fr.

L. leucothallina

[L. cinnabarina Somerf. |
(Ach.) Raes.

[Lecidea decipiens Ach.] =Psora decipiens Hoffm.
(Ignore the incorrect synonymy on the label)

L. dolodes Nyl. ex Hasse

L. elabens Fr.

L. elata Schaer.

Protoblastenia russula

[L. friesii Ach. in Liljebl.] =Hypocenomyce friesii

(Ach. in Liljebl.) G. Schneider
[L. icterica (Mont.) Tayl.] =Psora icterica (Mont.)
Muell.—Arg.
L. insularis Nyl.
L. leucothallina Arn.
L. limosa Ach.
[L. luridella Tuck. ]

L. luridella =Psora luridella (Tuck.) Fink
L. lyngei Degel.

Texas
Colorado
California

Colorado
Wyoming
California

Colorado
Colorado
California
Mexico
Colorado
Colorado
Galapagos

Colorado
Colorado

‘Texas

Colorado
Colorado
Australia

Colorado

‘Colorado

Oregon
Australia

Colorado
California
Australia
Colorado

Australia

N. Mexico
Montana
Montana
N. Guinea
Colorado
Colorado

436

451

273

155

428

272

435

424

[L. novomexicana (B. de Lesd.) W. A. Web. ex R. Ander—

son] =Psora nipponica (A. Zahlbr.) G. Schneider

L. nylanderi (Anzi) Th. Fr.

[L. quadricolor (Dicks.) Borr.] =L. granulosa
(Hoffm. ) Ach.

[L. rubiformis (Wahlenb. ex Ach.) Ach.] Psora tuck
ermanii R. Anderson ex G. Schneider, invalid. no
Latin description. __

[L. rubiformis (Wahlenb. ex Ach.) Ach.] Psora cere-
briformis W. A. Web., sp. nov.

[L. rufonigra (Tuck.) Nyl.] =Psorula rufonigra
(Tuck. ) G. Schneider

[L. symmicta (Ach.) Ach.] Lecanora cadubriae (Mass.)
Hedlund

[L. texana W. A. Web.] =Xanthopsora texana (W. A.
Web.) G. Schneider & W. A. Web. TYPE OOLL.

[L. wallrothii Flk. ex Spreng.] =Trapeliopsis wall-
rothii (Flk. ex Spreng.) H. Hertel & G. Schneider

Lecidella elaeochroma (Ach. ) Haszl.

[Leprocaulon albicans (Th. Fr.) Nyl. ex Hue]

rocaulon gracilescens M. Lamb & Ward

L. arbuscula (Nyl.) Nyl. ex Hue
[L. microscopicum (Vill.) Gams ex D. Hawksw. ]
L. tenellum (Tuck.) Nyl.
Leprocaulon tenellum (Tuck.) Nyl.
Leptogium brebissonii Mont. in Webb
L. burnetiae Dodge var. hirsutum (Sierk) P. M. Joerg.
L. cyanescens (Ach. ) Koerb.
L. denticulatum Nyl.
[L. foveolatum Nyl.] L. punctulatum Nyl.
[L. furfuraceum (Harm. ) Sierk]
Lesd. ) Dodge
L. hypotrachynum Muell-Arg.
L. javanicum (Mont. & v. d. Bosch) Mont.
L. lichenoides (L.) A. Zahlbr.

L. ml legranum Sierk

L. moluccanum (Pers. in Gaud.) Vain.
L. phyllocarpum (Pers.) Mont.
L. rugosum Sierk
SEES vulpina (L.) Hue
L. wulpina (L.) Hue f. californica (Lev.) W. A. Web.
Richina Lichina confinis (0. F. Muell.) C. A. Agardh

L. papillosum (B. de |

Idaho
Minnesota

Arizona

Colorado

Colorado
Texas

Colorado

Texas

California

Colorado

Arizona

Qolorado

N. Guinea

Galapagos
Peru
Australia
Colorado
Galapagos
Colorado
Mexico
Galapagos

Arizona
Australia
N. Guinea
Colorado
Oregon
Arizona
Arizona
N. Guinea
Mexico
Arizona

California

Nevada
Tasmania

224
536

103

124

L. tasmanica Henssen

Lobaria hallii Tuck.

L. retigera (Bory) Trevis.

[L. stictaeformis (Schaer.) Trevis.] L. discolor
(Bory in Del.) Hue

Lopadium pezizoideum (Ach. ) Koerb.

Maronella laricina Steiner ISOTYPES

Mastodia tessellata auct. (See Brodo, The Bryologist
79: 396-398. 1977, for nomenclatural discussion)

Melanaria melanospora (Nyl.) Erichsen

[Menegazzia aeneofusca (Muell.-Arg.) R. Sant.]
M. nothofagi P. James ined.

M. pertusa (Schrank) Stein

Micarea denigrata Fr.) Hedlund

Microthelia micula Koerb.

Mycoglaena myricae (Nyl.) R. C. Harris

Neophyllis melacarpa (F. Wils.) F. Wils.
G. Schneider maintains this over Yoshimura's
placement in Gymnoderma

Nephroma arcticum (L.) Torss.

N. australe A, Richard

N. cellulosum (Sm. ex Ach.) Ach.

N. helveticum Ach.

N. resupinatum (L.) Ach.

[Nephromopsis californica Gyel.] =Cetraria orbata
(Nyl.) Fink

Neuropogon acromelanus (Stirt.) M. Lamb

N. sulphureus (Koenig) Hellb.

Normandina pulchella (Borr.) Nyl.

Ocellularia alba (Fee) Muell.-Arg.

Ochrolechia grimmiae Lynge

O.upsaliensis (L.) Mass.

Omphalaria kansana Tuck. Determination doubtful.

Opegrapha saxicola Ach.

Pannaria leucophaea (Vahl) P. M. Joerg.

[P. nigrata Muell.-Arg.] Possibly P. tavaresii P. M.

Joerg., cf. Opera Bot. 45:70. 1978.

P. pezizoides (G. Web.) Trevis.

Parathelium sp. indet.

[Parmelia bostrychodes A, Zahlbr.] =Hypotrachyna
bostrychodes (A. Zahlbr.) Hale

[P. caperata (L.) Ach.] =Pseudoparmelia caperata
(L.) Hale

[P. chiricahuensis R. Anderson & W. A. Web. ]
=Neofuscelia chiricahuensis (R. Anderson & W. A.
Web.) Essl.

Tasmania
Oregon
India

N. Guinea
Idaho
Austria

Alaska
Chile

Australia

N. Guinea
Colorado
Arizona
Colorado

Australia
Alaska
Tasmania
Australia
Colorado
Alaska

California
Tasmania

Spitzbergen

Australia
Australia

Spitzbergen

Colorado
Colorado
California
USSR

Australia
Colorado
Chile

N. Guinea
Australia

Arizona
Texas

[P. conspersa (Ach.) Ach. ]
chroa (Tuck. ) Hale

[P. dichotoma Muell.-Arg. ]
toma (Muell.-Arg.) Hale

[P. dominicana Vain. ]
Hale

[P. elegantula (A. Zahlbr.) Szatala]
elegantula (A, Zahlbr.) Essl.

[P. endosulphurea (Hillm.) Hale]
sulphureun (Hillm. ) Hale

[P. exasperatula Nyl.] =Melanelia exasperatula
(Nyl.) Ess1.

[P. furfuracea (L.) Ach.] Pseudevernia intensa
(Nyl.) Hale & W. Culb.

[P. galbina Ach.] =Parmelina galbina (Ach.) Hale

[P. isidiotyla Nyl.] Neofuscelia loxodes (Nyl.)
Essl.

[P. novomexicana Gyel. ]
(Gyel.) Hale

[P. perlata (Huds.) Ach. ]
(Huds. ) Hale

[P. physodes (L.) Ach. ]
Nyl.

P. praesignis Nyl.

P. pseudotenuirima Gyel,

[P. pulla (Schreb.) Ach.] =Neofuscelia pulla (Ach.)
Essl.

[P. pulvinata Fee]
Hale

[P. revoluta Flk.] =Hypotrachyna revoluta (Flk.)
Hale

[P. rutidota Hook. f. & Tayl.] =Pseudoparmelia
rutidota (Hook. f. & Tayl.) Hale

P. saxatilis (L.) Ach.

[P. saximontana R. Anderson & W. A. Web.] TYPE
COLL. =Melanelia substygia (R. Anderson & W. A.
Web.) Essl.

P. signifera Nyl.

P. subalbicans Stirt.

[P. subcrinita Nyl.] =Parmotrema subtinctorium
(Nyl.) Hale

P. subrudecta Nyl.

Xanthoparmelia chloro-

=Xanthoparmelia dicho-

=Melanelia

=Parmotrema endo-

=Parmotrema perlatun

=Hypogymnia physodes (L.)

psy OL uae ayia pL nse S

[P. substrigosa Hale in W. A. Web.] =Xanthoparmelia

substrigosa (Hale in W. A. Web.) Hale
[P. subtinctoria A. Zahlbr.] =Parmotrema subtinc-

torium (A. Zahlbr.) Hale

=Parmotrema dominicana (Vain. )

=Xanthoparmelia novomexicana

Wyoming
Australia
Galapagos

Colorado

Australia
Colorado
Arizona
Australia
Colorado
N. Mexico
Australia
Australia
Arizona
Qolorado
Australia

Colorado

Colorado
Australia
Australia
Dominica
Australia

Australia

Australia
Arizona

[P. taractica Kremp.] =Xanthoparmelia taractica

(Kremp.) Hale
[P. tinctorum Nyl.] =Parmotrema tinctorum (Nyl.)
Hale

[P. ulcerosa A. Zahlbr.] =Xanthoparmelia ulcerosa
(A. Zahlbr.) Hale

[P. ulophyllodes (Vain.) Sav.] =P. soredica Nyl.

[P. weberi Hale] =Xanthoparmelia weberi (Hale) Hale

[P. wyomingica (Gyel.) Hale] =Xanthoparmelia wyo-
mingica (Gyel.) Hale

Parmeliopsis aleurites (Ach. ) Nyl.

P. placorodia (Ach.) Nyl.

Peccania sp. indet.
Peltigera aphthosa (L.) Willd.
Peltigera dolichorhiza (Nyl.) Nyl.

P. horizontalis (Huds.) Baumg.

P. horizontalis (Huds.) Baumg. f. zopfii (Gyel.)
J. W. Thomson
P. polydactyla (Neck. ) Hoffm.

[P. scabrosa Th. Fr.] P. sp. nov. Vitikainen ined.
P. venosa (L.) Willd.
Peltula cylindrica Wetmore
P. obscurans (Nyl.) Gyel. var. deserticola
(A. Zahlbr.) Wetmore
Pertusaria californica Dibben
P. saximontana Wetmore
P, xanthodes Muell.-Arg.
Pertusaria (Subg. Lecanorastrum) sp. indet.
[Phaeographina tridacna W. A. Web. sp. nov. ined.]
P. isidiosa (Vain.) A. Zahlbr., det. Nakanishi.
Phaeographis exaltata (Mont. & v. d. Bosch) Muell.-
Arg.

P. sp. indet.

Phylliscum demangeonii (Moug. ) Nyl.

Physcia adscendens (Th. Fr.) Oliv. em. Bitt.

[P. aegialita (Ach.) Nyl.] Dirinaria subconfluens
Awasthi, cited specimen. ©

P. caesia (Hoffm.) Hampe Identification incorrect,
but no alternative available.

P. callosa Nyl.

P. duplicorticata W. A. Web. & J. W. Thomson TYPE
COLL.

Arizona
Galapagos

Venezuela
Colorado
Arizona

Wyoming
Arizona
Colorado
Arizona
Mexico
Colorado
Australia
N. Guinea
Colorado
N. Guinea

Colorado
Alaska
Oregon
Colorado
Colorado
Georgia

Arizona

California

Colorado
Louisiana
Tasmania
N. Guinea
Mexico
Galapagos

Colorado
Colorado

Australia

Australia
Colorado

Calif.

[P. pulverulenta (Schreb.) Hampe f. coralloidea Suza
in Nadv.| Physconia sp. indet.

[P. setosa (Ach.) Nyl.] P. hispidula (Ach.) Frey

[P. setosa (Ach.) Nyl.] Identification incorrect.

P. stellaris (L.) Nyl.

P. tribacoides Nyl.

P. vitii Nadv.

Physconia detersa (Nyl.) Poelt

P. muscigena (Ach.) Poelt

Physma byrsinum (Ach. ) Muell.—Arg.

Pilophoron robustum Th. Fr.

Placynthium nigrum (Huds.) S. Gray

Platismatia stenophylla (Tuck.) W. Qulb. & C. Qulb.

Polychidium muscicola (Sw.) S. Gray

Porina epiphylla Fee

P. rubentior (Stirt.) Muell.—Arg.

P. sp. indet.

Pseudocyphellaria argyracea (Del. ) Vain.

P. australiensis H. Magn.

[P. durvillei (el.) Vain.] P. hirta (G. Forst.)
D. Gall. & P. James

[P. freycinetii (Del.) Malme] Identification dubious.

P. gilva (Ach.) Malme

P. glabra (Hook. f. & Tayl.) Dodge

P. impressa (Hook. f. & Tayl.) Vain.

P. neglecta (Muell.-Arg.) H. Magn.

[P. orygmaea (Ach.) Malme] P. hirta (G. Forst.)
D. Gall. & P. James

P. sp. indet.

P. rainierensis Imshaug

Psoroma soccatum R. Br. in Cromb.

P. sphinctrinum (Mont.) Nyl.

Psorotichia minuta H. Magn.

P. sp. indet.

Pyrenotrichum splitgerberi Mont.

Pyrenula cerina Eschw.

P. nitidella (Flk. ex Schaer.) Muell.-Arg.
var. maculata R. C. Harris TYPE OOLL.

Pyxine pringlei Imshaug

[Ramalina ceruchis (Ach.) DeNot.] =Desmazieria
ceruchis (Ach.) Trevis.

[R. combeoides Nyl.] =Desmazieria combeoides (Nyl.)
Follm. & Huneck

R. complanata (Sw. ex Ach.) Ach.

N. Mexico
Arizona
Australia
Colorado
Louisiana
Germany
Colorado
Colorado

Australia
Australia

Tasmania
Australia
Tasmania
N. Guinea
Tasmania
Tasmania
Australia

Tasmania
Australia

Washington

Australia
Tasmania
Colorado
Texas
Louisiana
Galapagos

Texas
Galapagos

Mexico

Mexico
Galapagos

R. farinacea (L.) Ach.

R. fraxinea (L.) Ach. Identification doubtful.

R. geniculata Hook. f. & Tayl. var. compacta Muell.—

Arg.
[R. homalea Ach.] =Desmazieria homalea (Ach. ) Mont.

R. javanica Nyl.

R. linearis (Sw.) Ach.

R. menziesii Tayl., non Tuck.
R. pacifica Asahina

R. peruviana Ach.

R. sinensis Jatta
R. sorediantha Nyl.

R. subleptocarpha Rundel & Bowler
R. usnea (1. ) Howe f.

[Ramalodium succulentum (R. Br.) Nyl.] Arctomia
fruticosa Henssen & Weber. sp. nov. ined.

Reinkella lirellina Darb.

R. parishii Hasse

Rhizocarpon macrosporum Raes.

Rinodina applanata H, Magn.

[R. archaea (Ach.) Vain. em. Malme] Sheard questions
the identification.

R. calculiformis W. A. Web. TYPE COLL.
R. coloradiana H. Magn.

R. svillensis H. Magn.

[R. oreina (Ach.) Mass.] =Dimelaena oreina (Ach.)
Norm.

[R. radiata (Tuck.) Tuck.] =Dimelaena radiata (Tuck.)
Hale & W. Culb.

R. turfacea (Wahlenb.) Koerb.

Roccella babingtonii Mont.

R. fimbriata Darb.

R. fucoides (Dicks.) Vain.

R. portentosa Darb.

[R. portentosa Darb.] R. galapagoensis Follm. TYPE
COLL.

Roccellaria mollis (Hampe) A. Zahlbr.

Roccellina condensata Darb.

Galapagos
California
Australia

Tasmania
Mexico
California

Philippines
Galapagos
N. Mexico
Galapagos
Galapagos
Mexico
Panama
Galapagos

Tasmania
Peru
California
Colorado
Louisiana

California -

California
Mexico
Colorado
California

Colorado

California
Colorado
Mexico
Galapagos
Mexico
Mexico
Italy
Chile

Galapagos
Galapagos
Chile
Peru
Chile

[R. lutosa Follm.] Roccellina luteola Follm.
Sarcogyne clavus (Ram. ) Kremp.

S. regularis Koerb.

Schismatomma cupressum Herre

Schistophoron tenue Stirt.

Schizopelte californica Th. Fr.

Siphula coriacea Tayl. ex Nyl.

Siphula fragilis (Hook. f. & Tayl.) J. Murray, nom.
nud, x ria

S. sp. indet.?

Speerschneidera euploca (Tuck. ) Trevis.

Sphaerophorus melanocarpus (Sw. ) DC.

S. tener Laur.

Sporastatia testudinea (Ach.) Mass.

Staurothele clopima (Wahlenb. ex Ach.) Th. Fr.

Stereocaulon glareosum (Sav.) H. Magn.

S. leprocauloides M. Lamb TYPE OOLL.
S. massartianum Hue

[S. microcarpum Muell.-Arg. ]
TYPE OOLL.

S. myriocarpum Th. Fr.

[Stereocaulon piliferum Th. Fr.] misspelling for
S. piluliferun th. Fr.

S. pseudomassartianum M. Lamb ex Frey

S. weberi M. Lamb,

S. ramilosum (Sw.) Raeusch var. nudatum (Muel1.-Arg.)
Muell.—Arg.

S. rivulorum H. Magn.

S. staufferi M. Lamb ex Frey

S. virgatum Ach. ex Spreng.

Sticta boschiana Mont. & v. d. Bosch ex Jungh.

S. filix (Sw.) Nyl. ait

S. [Pseudocyphellaria] cf. fragillima Bab. ex Hook.,
sensu A. Zahlbr.

S. fuliginosa (Dicks. ) Ach.

S. rubella Hook. f. & Tayl.

-S. weigelii Isert. ex Ach.

Strigula elegans (Fee) Muell.-Arg.

Teloschistes exilis (Michx.) Vain.
T. fasciculatus Hillm.
T. flavicans (Sw.) Norm.

T. velifer F. Wils.

Chile
Texas
Texas
California
Galapagos
Califormia
Australia
Australia

Tasmania
California
Texas
Tasmania
Tasmania
Colorado
California
Colorado
Montana

N. Guinea
N. Guinea

Philippines

Galapagos
N. Guinea

India

N. Guinea
N. Guinea
N. Guinea

Australia
Colorado
N. Guinea
Dominica
N. Guinea
Australia

N. Guinea
Australia
Tasmania
Arizona
Louisiana
Australia
Texas
Australia
Galapagos
Australia

100

T. villosus (Ach. ) Nor.
Thamolia vermicularis (Sw.) Schaer.

T. vermicularis (Sw.) Ach. ssp. solida (Sato)
W. A. Web.

[Thelidea corrugata Hue] =Knightiella splachnirima

(Hook. f. & Tayl.) Gyel.
Thelotrema diminitum Hale, Cited specimen.
Thrombium epigaeum (Pers.) Wallr.
Thysanothecium hookeri Berk. & Mont.
T. hyalinum (Tayl.) Nyl.
Toninia bullata (Mey. & Flot.) A. Zahlbr.
[T. ruginosa (Tuck.) Herre] T. sp. nov. ined.
Teetristiseinesrr,
Trypethelium grossum Muel1.-Arg.
T. mastoideum (Ach.) Ach.
T. tropicum (Ach.) Muell.-Arg.
Umbilicaria arctica (Ach.) Nyl.
U. coriacea Imshaug
U. cylindrica (L.) Del. ex Duby
U. decussata (Vill.) A. Zahlbr.
U. deusta (Huds.) Baume.
. havaasii Llano
. hirsuta! (Sw. ) Ach.
. hyperborea (Ach.) Hof fm.
. kraschenimnikovii (Sav.) A. Zahlbr.
. phaea Tuck.
. proboscidea (L.) Schrad.
subglabra (Nyl.) Harm.
U. vellea (L.) Ach.

qaqa, a qj, ca

[U. vellea (L.) Ach.] U. cinereorufescens (Schaer.)

Frey sensu Llano
Usnea angulata Ach.
U. angulosa (Muell.—Arg.) Mot.
U. arbusculiformis Mot.
U. capillacea Mot.

U. cavernosa Tuck. in Agassiz

U. cf. cladocarpa Fee
U. comosa (Ach.) Roehl.

[U. diplotypus Vain.] U. herrei Hale. nom. nud.

U. aff. elongata Mot.
U. flexilis Stirt.

Mexico
Colorado
Alaska
Australia

N. Guinea

Tasmania
Australia
Colorado
Australia
Australia
Australia
Colorado
Colorado
N. Guinea
Louisiana
Louisiana
Norway
Norway
Australia
Australia
Colorado
Norway
Colorado
Colorado
Colorado
California
Australia
Australia
Colorado

Colorado
Mexico
Australia
Mexico
Tasmania
Tasmania
Arizona
Mexico
Galapagos
Arizona
Kansas
Texas
Galapagos
Tasmania
N. Guinea

. herrei Hale, nom. mud.
. 1gniaria Mot.
. inermis Mot.
. cf. merrillii Mot.
. microcarpoides (Muell.-Arg.) Mot.
. mutabilis Stirt.
. paradoxa (A. Zahlbr.) Mot.
. poliotrix Kremp.

aIaI1al acl aiciacjia

[U. rubiginea (Michx.) Mass.] U. rubicunda Stirt.
U. scabrata Nyl. ssp. nylanderiana Mot.
U. scabrida Tayl.

[U. sorediifera auctt.] =U. fulvoreagens (Raes.) Raes.

cf. Brodo in The Bryologist 79:406. 1976 [1977].

U. spilota Stirt.
U. cf. strigosa (Ach.) Eaton
U. torquescens Stirt.

U. torulosa (Muell.-Arg.) A. Zahlbr.

[U. tristis Mot.] U. florida (L.) Wigg., fide
Asahina, J. Jap. Bot. 43:65. 1968. eae

U. xanthopoga Nyl.

Verrucaria laevata Ach.

‘Xanthoparmelia wyomingica (Gyel.) Hale

Xanthoria candelaria (L.) Th. fr.
X. fallax (Hepp ex Arn.) Arn. sens. latiss.
X. polycarpa (Ehrh.) Rieb.

X. polycarpa (Ehrh.) Rieb. Identification doubtful.

[Xylographa spilomatica (Anzi) Th. Fr.] =X. vitiligo

Ach. ) Laundon
X. vitiligo (Ach. ) Laundon

NEW COMBINATIONS

Cladina galapagosensis (Ahti) W. A. Web.
Ahti, Ann. Bot. Soc. 'Vanamo' 32(1):46. 1961.

Cladina polia (R. Sant.) W. A. Web.
orth. error) R. Sant., Ark. Bot. 30A(10): 15. 1952.

Heterodemmia barbifera (Nyl.) W. A. Web.
syn, Lich. 1:416. 1860.

Heterodemmia circinalis (A. Zahlbr.) W. A. Web.

Arizona
Chile
Tasmania
Mexico
Australia
Texas
Galapagos
Australia
Australia
Australia
Australia
USSR
Tasmania
Tasmania

Colorado
Colorado
Wyoming
Australia
Mexico
Australia
Australia
Tasmania

N. Mexico
Tasmania
S. Dakota
Colorado
Mexico
Texas
Colorado
California

Colorado
Montana

Cladonia galapagosensis
Cladonia polia (as "pohlia”,
Physcia barbifera Nyl.,

Anaptychia leuco-

melaena var. mltifida f. circinalis A. Zahlbr., Beih. Bot. Centralbl.

19(2):84. 1905.

102

Heterodermia stellata (Vain.) W. A. Web. Anaptychia podocarpa
var. stellata Vain., Acta Soc. Fl. Faun. Fenn. 7:131. 1890.
Heterodemnia verrucifera (Kurokawa) W. A. Web. Anaptychia leucome-
laena f. verrucifera Kurokawa, Nova Hedwigia, Beih. 6:72. 1962.
VALIDATIONS OF NEW SPECTES

LECANORA PSEUDOPINGUIS W. A. Web., sp. nov. Thallus saxicolous crus-
taceus effusus indeterminatus rimoso-areolatus, areolis minus quam 1 pm
diam ca. 0.5 mm crassis laevibus pallide citrinis vel albescentibus so-
redia et isidia desunt. Cortex 30m crassus, granulis lutescentibus in-
spersis, medulla fulvescens, C+ aurantiaca, P-, K-, IKI-, UV+ rubro-viol-
ascens. Apothecia sessilia usque ad 1.5 mm diam margine crasso laevi vel
undulato-crenulato disco plano livido pulverulento, epihymenio flavo-gra-
nuloso C+ aurantiaco hymenio 50-60 jam crasso sporis 8/nae 7-9 x 3-5 um.
Pycnidia cylindrica 200Mm alta x 100 sm lata margine prominenti elevato,
conidiis arcuatis acicularis 15-20 ym longis. Ad saxa wulcanica praeci-
pue litoralis.

Thallus saxicolous, crustaceous, effuse, indeterminate, irregularly ri-
mose-areolate with areoles less than | mm diam, ca. 0.5 mm thick, smooth,
pale citrine to albescent, lacking soredia or isidia. Cortex 30 pm
thick, inspersed with yellowish granules; medulla discolored, C+ orange,
P-, K-, IKI-, UV+ red-violet. Apothecia sessile, up to 1.5 mm diam, with
thick smooth or crenulate-wavy margin, the disk plane, livid, pulveru-
lent; epihymenium with yellow granules, C+ orange; hymenium 50-60 Mm
high, spores 8/ascus, 7-9 x 3-5 44m; pycnidia cylindric, 200 jim deep x 10
yim wide, with prominent raised rim resembling the apothecial margin; pyc-
noconidia arcuate, acicular, 15-20141m long. On volcanic rocks, primarily
littoral.

HOLOTYPE. Ecuador. Galapagos Islands. Isla Santa Cruz, vicinity of
Academy Bay, on exposed point along shore just east of Darwin Research
Station, abundant on rocks just above high tide mark, 10 April 1976,
Weber & Lanier, Lich. Exsicc. OOLO No. 500 (COLO). Further distribu-
tion. Floreana I., Jervis I., Wenman I., S. Plazas Islet, Champion I.,
and probably on every island of the archipelago. One population has been
found inland from the seacoast but on a seaward exposure: Santiago I., on
rocks at rim of easternmost crater in the highlands, 400 msm, 10 May
1971, L. H. Pike 2112 (COLO L-55204). This is a most unusual occurrence
and may be correlated with atmospheric conditions (upwelling of fog) pe-
culiar to the site. L. pseudopinguis is one of the dominant species co-
loring the rocks of the shoreline yellow on many of the islands.

_ Except for its effuse thallus the new species resembles the lobate
littoral Californian Lecanora pinguis Tuck. Dr. Chicita Culberson, to
whom I am indebted for analyses of the chemistry of L. pseudopinguis and

103

the next species, reported: "'L. pseudopinguis contained a compound (Re
classes A4-5, B5, C4-5; spot color with 10% H»S0, and heat is orange
brown) in high proportion that is probably a xanthone pigment....We have
seen this same pigment before in a sample from Ahmadjian that was also
from the Galapagos Islands but that had been identified as Buellia stra-
minea. The proportions of the minor products of Ahmadjian's sample were
different from the Lecanora, and the Lecanora also contained a trace of
thiophaninic acid and another unknow pigment (probably not a xanthone)
that is common in lichens and that we have previously called "pigment
SV-1".

LECANORA (Eulecanora) TEXANA W. A. Web., sp. nov. Prothallus
destitutus. Thallus effusus indeterminatus laevis vel tartareus non- vel
indistinctissime lobatus, usque ad 2 mm crassus, irregulariter rimoso-
areolatus vel continuus, pallide flavus. Pseudocortex prosenchymaticus
30-40 41m crassus ex hyphis erectis, strato algarum 30-50 yam alto, con-
tinuo. Medulla densissima, alba, subtus fulvescens, K-, KC+ luteus.
Apothecium immersum vel adnatum, usque ad 2 mm diam, margine vestigiali,
excipulo proprio ex hyphis erectis parallelis, disco carneo, plano vel
convexiusculo, epihymenio hyalino granuloso, granulis in K dissolvens,
hymenio 50-60 jAm alto, subhymenio ex hyphis verticalibus, hypothecio hya-
lino subtus algifero, ascis IKI+ coeruleis, paraphysibus simplicibus, 2
m diam, gracilibus, non capitatis, sporis 8-nae anguste-ellipsoideis
rectis vel plus minusve curvatis, 10-14 x 4-5 wm. Pycnidium ostiolo
aeruginoso loculo 200 pm diam, pycnoconidiis Eulecanoroideis acicularis,
arcte curvatis vel sigmideis, 10-15 x 0.5 jam. Ad saxa granitica, areas
amplas rupium superficieram verticalibus tectus.

Prothallus lacking. Thallus effuse, indeterminate, smooth to tar-
tareous, very distinctly if at all marginally lobate, up to 2 mm thick,
irregularly rimose-areolate or continuous, pale yellow; pseudocortex pro-
senchymatous, 30-40 ym thick, of erect hyphae; algal layer 30-50 m high,
continuous; medulla very dense, white, becoming discolored brownish be-
low, K-. KC+ yellow wash. Apothecia immersed to adnate, up to 2 m dian,
the margin vestigial or represented by a few small bead-like remnants;
proper exciple of erect parallel hyphae; disk flesh-colored, plane to
slightly convex; epihymenium hyaline, granulose, the granules dissolving
in K; hymenium 50-60 pm high, difficult to differentiate fram subhymenium
of vertical hyphae; hypothecium hyaline, with continuous or broken algal
layer below; asci IKI+ blue; paraphyses simple, 2 jAm diam, slender, not
capitate; spores 8/ascus, narrowly ellipsoid, straight or somewhat
curved, 10-14 x 4-5qm. Pycnidia appearing as black dots on thallus sur-
face, the ostiolar margin aeruginose, the loculus 200 Mm diam; pycno-
conidia of Fulecanora type, acicular, curved in a semicircle, 10-15 x 0.5
fam. On granite rocks, covering large areas of the vertical faces, rarely
fruiting, but when fertile with clusters of apothecia in small plaques of
the thallus.

104

HOLOTYPE. Texas, U. S. A. Gillespie Co. [not Llano as given on the
label]: trail to 'Balanced Rock'"', a granite dome rising above the sur-
rounding plain, 4 mi n of Fredericksburg in the Texas "Hill Country";
abundant and dominant on vertical N-facing massive granite blocks, 29
April 1974, Weber, Lich. Exsicc. COLO No. 451 (COLO).

Lecanora texana is apparently closely related to L. sulphurea
(Hoffm.) Ach., but the apothecium is pale from the beginning, never
blackening. Dr. C. Culberson reported: "L. texana contained usnic acid,
zeorin and a trace of atranorin. In addition there were two or three un-
identified triterpenoids that are probably related to zeorin but which do
not correspond to leucotylin or leucotylic acid (the only known compounds
of this type in our collection). There is also a curious unidentified
phenolic compound (R¢ classes A3, B2-3, C5; spot color with 10% H»SO, and
heat is yellow). This may be a new lichen product."

PSORA CEREBRIFORMIS W. A. Web., sp. nov. Thallus terricola creta-
ceus tumulos hygroscopicos usque ad 5 cm diam formantes, squamilis dense
aggregatis sed non imbricatis primo planis mox alte convexis rimosis mar—
ginibus revolutis. Cortex argillaceus pseudoparenchymaticus 50-60 m
crassus, strato epinecroso crasso albo tectus. Stratum gonidialium con-
tinuum. Medulla alba solida crassa, coacto rhizo-hyphoso substrato af-
fixo. Cortex et medulla K-, C-, KC-, P-, IKI-. Apothecia 1-2 m diam
atra marginalia convexa numerosa plerumque aggregata, excipulo ex hyphis
parallelis pallide ferrugineo, ascis clavatis 30-40 m longis, sporis el-
lipsoideis (8)10-12(17) x 5-6 m. Epihymenium ferrugineum K+ vinosum C-,
Pp. IKI.

Thallus terricolous, chalk-white, forming hygroscopic convex mounds
up to 5 cm or more diam; squamles densely aggregated but not overlapp-
ing, up to 5 mm diam, at first plane but very soon highly convex, deeply
cracked, with the margins rolled under. Cortex pale clay-colored, of
pseudoparenchyma type, 50-60 m thick, covered by a thick white epinecral
pruinose layer. Algal layer continuous. Medulla white, solid, thick,
attached to the substrate by a rhizo-hyphal felt. Reactions K-, C-.
KC-. P-. IKI-. Apothecia 1-2 mm diam, black, marginal, highly convex,
abundant and often clustered; exciple of parallel hyphae, pale reddish-
brown; asci clavate,30-40 m long; spores ellipsoid, (8)10-12(17) x 5-6
m; epihymenium reddish-brown, K+ vinose, C-, P-, IKI-.

HOLOTYPE. U. S. A. Colorado. Montrose Co.: on gypsum knolls, floor
of Paradox Valley 4 mi E of Bedrock, 1,500 msm, 30 May 1960, Shushan,
Anderson & Weber, Lich. Exsicc. COLO No. 24 (COLO).

MYCOTAXON

WO lee AIL ORmeL ee Oye meio LY April-June 1981

CULTURAL STUDIES ON PORIA CINERASCENS,,
P. RIVULOSA, AND P. SUBVERMISPORA

(Aphyllophorales, Basidiomycotina)

K. K. NAKASONE

Center for Forest Mycology Research
1/

Forest Products Laboratory—
USDA, Forest Service
Madison, oWLs 5 05/05

ABSTRACT

The culture complex involving Poria cinerascens, P. rivulosa,
and P. subvermispora is examined. Cultural descriptions are provided
for each species. Although nearly identical, cultures of these three
species can be separated by temperature studies, growth on cabbage
extract agar, and microscopic characteristics.

INTRODUCTION

Sporophores of Poria cinerascens (Bres.) Sacc. et Syd., Poria
rivulosa (Berk. et Curt.) Cke., and Poria subvermispora Pilat are dis-
tinct and easily identified by observing the hyphal systems and basid-
iospores (Lowe 1966). However, when grown in pure culture they are
nearly indistinguishable. Nobles (1948) treated P. rivulosa (as P.
albipellucida Baxter) and P. cinerascens together under one culture
description. Later, Nobles (1965) and Stalpers (1978) distinguished
these fungi on minor or variable characters.

Because of the difficulty in identifying these cultures, the im-
portance of individual species of the complex in the decay of products,
preservative-treated wood, and other wood forms cannot be assessed
accurately. Duncan and Lombard (1965), in their summary report on
Hymenomycetes associated with decay of wood products, report these
fungi collectively as the Poria cinerascens complex. Thus, this study
was undertaken to resolve the P. cinerascens culture complex.

MATERIALS AND METHODS

Cultures were grown at 25° C in the dark on 1.5% malt extract agar
(MEA) in glass petri plates and were examined at weekly intervals.

1/ Maintained at Madison, Wis., in cooperation with the University
of Wisconsin.

106

Cultures were grown also on 0.5% gallic acid agar (GAA), on 0.5% tan-
nic acid agar (TAA) (Davidson et al., 1938), and on cabbage extract
agar (CAB) at 25° C and measured at 7 and 14 days.

Cabbage extract agar was made from fresh cabbage because growth |
of these fungi on commercial media was not satisfactory. CAB is pre-
pared as follows: Place 500 gm chopped, fresh cabbage in 1000 ml dis-
tilled water, cover, and bring to a boil. Simmer about 15 hours (over
several days if necessary) and add about 1,000 ml distilled water
during this time. Filter through cheese cloth and bring final volume
of cabbage extract to 1000 ml. Add 20 gm agar, 2.5 gm diabasic sodium
phosphate, 5 gm sodium chloride, and 10 gm Difco Bacto-peptone to the
1000 ml of cabbage extract and autoclave.

Unless otherwise indicated, all cultures studied are of polyspo-
rous origin. Monosporous isolates were obtained from fruiting in cul-
ture. Key patterns of 2-week-old cultures follow the format of
Davidson et al. (1942). Species codes of 6-week-old cultures follow
the format of Nobles (1965). Cultures are on deposit at the Center
for Forest Mycology Research.

Sporophores associated with the cultures were previously identi-
fied by Drs] J: L. Lowe, Ri L. Gilbertson, or) M.J.,3Llarsen. 7 sAL though
the sporophores were not reexamined by the author, dikaryotic-
monokaryotic matings were done with all cultures studied. Cultures
listed under each species had successful matings with one or more hap-
loids of that species.

CULTURE DESCRIPTIONS

Poria Cinerascens (Bres.) Sacc. et Syd., Syil. Fung. 1671615 81902"

Growth characters: Growth on MEA rapid, plates covered in 1 wk;
mats white, appressed, thin, subfelty, with a network of thin, radial
strands, sometimes slightly raised and downy or felty at 2 wk, may |
develop small white balls of mycelium at 6 wk; margin even, appressed;
odor none; agar discoloration none; not fruiting by 6 wk. Oxidase
reactions at 1 wk on GAA moderately strong, mat 37-51 mm diam; on TAA
strong, mat 12-26 mm diam. Optimum temperature 36° C (Fig. 1). Growth
on CAB O-trace at 1 wk.

Microscopic characters: Hyphae of advancing zone 6-8 wm diam,
thin-walled, simple septate, branched, becoming thick-walled and scat-
tered by 2 wk. Hyphae of submerged mycelium 2-4 wm diam, thin-walled,
nodose septate, branched. Hyphae of aerial mat (a) similar to sub-
merged hyphae except occasionally encrusted with hyaline crystals; (b)
"binding hyphae" very slender, 0.5 um diam, aseptate, richly branched,
hyaline, nonstaining in phloxine, rare to abundant at 2 wk; (c) fiber
hyphae (Fig. 2) 4-6 um diam, thick-walled, walls thinning toward apex,
aseptate, hyaline nonstaining in phloxine, apparently lacking or numer-
ous at 2 wk. Chlamydospores globose to limoniform, 8-13.5 x 8-13.5 um,
thin-walled, walls thickening in age, hyaline, terminal or intercalary,
few to abundant in aerial and submerged mycelium. Bulbils sometimes
present in submerged mycelium.

Key patterns: A-P-F-1-2-10; A-P-F-1-2-10-16; A-P-F-1-2-11; A-P-F-
1=2-11-16; A-P-F-1-2-10-14.

Species code?) 2740072 (8 )) (22 494.6238, 42-55 250.

Monosporous cultures: Eight monosporous cultures of HHB 76 (4 of
each mating type) examined were similar to the polysporous cultures
except that they did not have clamp connections.

PORIA SUBVERM/ISPORA

RADIUS (mm)
S

20
PORIA RIVULOSA

PORIA CINERASCENS

/6 20 24 28 32 36 40 44
TEMPERATURE (°C)

Fig. 1. Average radial growth of Poria species on MEA
after 4 days at 8 temperatures. (M 149 135)

10 um

M 149 140

Fig. 2. Fiber hyphae from a culture of Poria cinerascens

(FP 100506) at 2 wk.

Fig. 3. "Binding hyphae" from cultures of Poria rivulosa

and P. subvermispora at 2 wk. (M 149 140)

107

108

Incompatibility system: Poria cinerascens has been reported by
Nobles et al. (1957) to be heterothallic and bipolar. Pairing of |
17 monosporous cultures of HHB 76 confirm their finding: Ay =01,45,0.,
Sells. Osos A, Se Oe Onis Pe Ole slates

Cultural description: Nobles (1948); Singh (1966).

Cultures studied: U.S.A.: MARYLAND--FP 100506 on pine,

Prince Georges County; SOUTH DAKOTA--FP 105939 on Pinus ponderosa
Dougl. ex Laws: (ponderosa pine), Black Hills National Forest, TEXAS--
HHB 76 cn Pinus sp., San Jacinto County; WISCONSIN--FP 105349 on Pinus
banksiana Lamb. (jack pine), Wood County.

Remarks: The development of fiber hyphae and optimum temperature
of 36° C are diagnostic for P. cinerascens. Poria cinerascems can
quickly be separated from P. rivulosa and P. subvermispora by growing
the cultures on MEA at 40° C (Fig. 1). At 4 days P. cinerascens grows
significantly more than the other species. Cultures of P. cinerascens
will often deteriorate after growing several years on artificial media.
Deteriorated cultures are slow growing, sodden, and do not develop
fiber hyphae.

Poria cinerascens is widely distributed in the U.S.A. and is
associated with a white rot of conifers.

Poria rivulosa (Berk. et Curt.) Cke., Grevillea 14:109. 1886.

Growth characters: Growth on MEA rapid, plates covered in 1 wk;
mats white, azonate, at 1 wk appressed, thin, subfelty, at 2 wk
slightly raised, thicker, subfelty to felty; margin even, appressed;
odor none; agar discoloration none; not fruiting by 6 wk. Oxidase
reactions at 1 wk on GAA moderate, mat 64-90 mm diam; on TAA strong,
mat 19-29 mm diam. Optimum temperature about 32° C (Fig. 1). Growth
on CAB 20-48 mm diam at 1 wk.

Microscopic characters: Hyphae of advancing zone 6-7.5 mm diam
thin-walled, simple septate, branched, by 2 wk often becoming thick-
walled, rare to abundant, absent in older cultures. Hyphae of sub-
merged mycelium 3-5 mm diam, thin-walled, nodose septate, branched.
Hyphae of aerial mat (a) 1.5-5 mm diam, similar to submerged hyphae,
sometimes encrusted with hyaline crystals; (b) "binding hyphae"

(Fig. 3) very slender, 0.5 mm diam, aseptate, richly branched, hyaline,
nonstaining in phloxine, usually present by 2 wk in clusters, difficult
to separate. Chlamydospores globose to limoniform, 8.5-11.5 x

8.5-11.5 mm, thin-walled, becoming thick-walled in age, hyaline, termi-
nal or intercalary, abundant in aerial mat.

Key patterns: -A-P-F-1-2-10-16; A-P-F-1-2-10; A-P-F-1-2-10-14-16;
A>P-F=1]=2-L0=14%

Species code: 2.4.(7).(8).34.36.38.41.42.55.59.

Monosporous cultures: Eight monosporous cultures of FP 133035 (4
of each mating type) were similar to the polysporous cultures except
that thick-walled advancing zone hyphae were present in 6-wk-old cul-
tures, and they lacked clamp connections and "binding hyphae."

Incompatibility system: Nobles et al. (1957) reported that P.
rivulosa is heterothallic and bipolar. Pairings of 17 monosporous Cute
tures of FP 133035 confirm their result: A) =01,35455, 9.12 15 LS,

i A, 2 05 7.,6,.0. bilan

Cultural descriptions: Buckland (1946), Baxter (1937), and Nobles
(1948) as Poria albipellucida Baxter; Stalpers (1978) as P. lindbladii
(Berk. et Br. ex Berk.) Cooke.

109

Cultures studied: U.S.A.: CALIFORNIA--Piirto 26 (rot) in
Sequoiadendron giganteum (Lindl.) Bucholz (giant sequoia); JLL 10602
on Sequoia sempervirens (D. Don) Endl. (redwood), Del Norte County;
FP 104207 (rot) in redwood, Wilson Creek; IDAHO--JLL 6987 on conifer,
Kootenai County; OREGON--FP 133035 (rot) in Tsuga sp., Benton County,
FP 133406 and FP 133696 on conifer, Lincoln County. CANADA:

BRITISH COLUMBIA--DAOM 9757 on Pseudotsuga menziesii (Mirb.) Franco
(Douglas-fir).

Remarks: Poria rivulosa is most similar culturally to P.
subvermispora. They can be separated, however, by growing the cul-
tures on CAB. Poria rivulosa will grow 20-40 mm in diameter at 1 wk
but P. subvermispora only O-trace.

Poria rivulosa is common in the northwest U.S.A. and
British Columbia on conifers. It is associated with a white ring rot
of living western redcedar, Thuja plicata Donn ex D. Don (Buckland
1946) and living redwood (Kimmey and Lightle 1955).

Poria subvermispora Pilacesstud iam Bote Cech 5:26 L94Gs

Growth characters: Growth on MEA rapid, plates covered in 1 wk;
mats white, azonate, at 1 wk appressed and subfelty or raised and
downy, at 2 wk appressed, subfelty to felty, or raised and woolly,
sometimes tufted; margin even, appressed; odor none; agar slightly
bleached in age; not fruiting by 6 wk. Oxidase reactions at 1 wk on
GAA moderate, mat 69-90 mm diam; on TAA strong, mat 22-30 mm diam.
Optimum temperature 32° C (Fig. 1). Growth on CAB O-trace at 1 wk.

Microscopic characters: Hyphae of advancing zone 6-7 wm diam,
thin-walled, simple septate, infrequently branched, by 2 wk becoming
thick-walled, rare. Hyphae of submerged mycelium 2-4 wm diam, thin-
walled or with slight wall thickening, nodose septate, branched.
Hyphae of aerial mat (a) similar to submerged hyphae, occasionally
encrusted with hyaline crystals; (b) "binding hyphae" very slender
(Fig. 3), 0.5 wm diam, aseptate, richly branched, hyaline nonstaining
in phloxine, usually present in clusters by 2 wk, difficult to sepa-
rate; (c) fiber hyphae rare, 2 wm diam, thick-walled, aseptate, hya-
line nonstaining in phloxine, infrequently branched, seen only in
6-wk-old cultures of FP 90031. Chlamydospores globose to limoniform,
1125-13.5 (-16) x 7.5-13.5 (-16) um, thin-walled, walls thickening in
age, hyaline, terminal or intercalary, numerous in submerged and
aerial mycelium.

Key patterns: A-P-F-1-2-10-16; A-P-F-1-2-10; A-P-F-1-2-10-14-16;
eesti 2—10=14.

Species codes.) 2-4, (7)-(8).34.36.38.41.(42).54.55.59.

Monosporous cultures: An examination of 8 monosporous cultures
obtained from JLL 14807 (4 isolates of each mating type) reveals that
they are similar to the polysporous isolates except that their mats
tend to be appressed, develop delicate radial to reticulate patterns,
and lack clamp connections.

Incompatibility system: Nobles et al. (1957), as P. notata
Overh., and Domanski (1969), as Fibuloporia subvermispora (Pilat)
Doman., reported that P. subvermispora is bipolar. Pairings of
17 monosporous cultures of JLL 14807 confirm their results: AL 1 ae
Bros ol2, 13,15, 16,18 519; A, Seow Ln lis Lee La

Cultural descriptions: Baxter (1947) (as P. quercuum Baxter);
Nobles (1948); Domanski (1969); Stalpers (1978).

110

Cultures studied: U.S.A.: ARIZONA--JLL 8904 on ponderosa pine,
Cochise County; COLORADO--JLL 6133 on conifer, Garfield County;

JLL 6332 on lodgepole pine, Larimer County; FLORIDA--FP 104027 (sporo-
phore tissue) on Quercus laurifolia Michx. (laurel oak), Duval County;
MARYLAND--FP 90031 on oak and FP 105752 on hardwood board,

Prince Georges County; NEW YORK--JLL 3292 on conifer, Essex County;
JLL 15225 on hardwood, Ulster County; OREGON--ME 485 (rot) in Douglas-
fir wood chips, Clatsop County. CANADA: Saskatchewan--JLL 14807 on
Populus tremuloides Michx. (aspen).

Remarks: Poria subvermispora is widely distributed throughout
continental U.S.A. It is associated with a white rot of conifers and
hardwoods and has been isolated several times from wood chip piles in
Washington and Oregon.

DISCUSSION

Cultures of P. cinerascens, P. rivulosa, and P. subvermispora can
be identified by noting several critical characters. These are sum-
marized in Table 1. Dikaryotic-monokaryotic pairings can be used also
to identify cultures once monosporous cultures are obtained. Cultures
of the three species will fruit on MEA and discharge spores if kept
long enough. The shape and size of the basidiospores can also be used
to differentiate between the species (Table 1).

Development of the slender "binding hyphae" found in the aerial
mat of all three species has not been recorded previously. Their
development in culture suggests their presence in the sporophore; how-
ever, this is not the case. Poria rivulosa and P. subvermispora have
monomitic hyphal systems (Lowe 1966). However, Poria cinerascens spo-
rophores have a trimitic hyphal system (Lowe 1966), and this is sup-
ported in culture.

Table 1.--Characteristics that differentiate cultures of
Poria cinerascens, P. rivulosa, and P. subvermispora

Optimum Growth Growth BE taal
growth at 1/ ony Host ate
temperature 40° C-— CAB-— size—
Ree oi ae REE ed thin ae my mm; ?s ero as ym
BOEEG 36 To ela Gp Oo
Hane rec eee - -trace Conifers
aaa GUAR ee Eee 1. Ser 220
5S 0e="6.0
P. rivulosa SZ La 20 - 48 Conifers me
4, Oo=e 500

Conifers 3250 ORO
P. subvermispora OZ Tae BO-L race and x
hardwoods 1.0 peed ico

1/ Average radial growth after 4 days on malt extract agar.
2/ Average mat diameters on cabbage extract agar after 7 days
ate 5o Se

3/ Spore measurements taken from Lowe (1966).

yal

Culturally, these three species belong to a unique group of fungi.
The rapid growth, production of extracellular oxidases, simple septate
hyphae of the advancing zone that later give rise to nodose septate
hyphae, and the bipolar mating system are characters shared by all spe-
cies of Group 54 (Nobles 1958). Phlebia subserialis (Bourd. et Galz.)
Donk, Phlebia subochracea (Bres.) Erikss. et Ryv., and Hyphodontia
setulosa (Berk. et Curt.) Maas G. are the most culturally similar to
the Poria species discussed.

ACKNOWLEDGMENTS

Ms. Ruth G. Payne is responsible for the successful development
and application of the cabbage extract medium. Critical reviews of
this manuscript, by Drs.) Row. Gilbertson, H.-H. Burdsall, Jr.=, and
Mrs. F. F. Lombard are greatly appreciated.

LITERATURE CITED

BAXTER, D. V. 1937. Some resupinate polypores from the region of the
Great Lakes. IX. Pap. Michigan Acad. Sci. 23:285-305, (published in
1938). BAXTER, D. V. 1947. Some resupinate polypores from the region
of the Great Lakes. XIX. Pap. Michigan Acad. Sci. 33:9-30, (published
in 1949). BUCKLAND, D. C. 1946. Investigations of decay in western
red cedar in British Columbia. Can. J. Res., C., 24:158-181.

DAVIDSON, R. W., W. A. CAMPBELL, and D. J. BLAISDELL. 1938. Differ-
ention of wood-decaying fungi by their reactions on gallic or tannic
acid medium. J. Agric. Res. 57:683-695. DAVIDSON, R. W.,

W. A. CAMPBELL, and D. B. VAUGHAN. 1942. Fungi causing decay of
living oaks in the eastern United States and their cultural identifi-
cation. USDA Tech. Bull. 785, 65 p. DOMANSKI, S. 1969. Wood-
inhabiting fungi of Bialowieza virgin forest in Poland. X. Fibuloporia
subvermispora (Pilat) Doman. comb. nov. and its diagnose. Acta Soc.
Bot. Pol. 39:531-538. DUNCAN, C. G., and F. F. LOMBARD. 1965. Fungi
associated with principal decays in wood products in the United States.
eos POr.e serv, Res. Pap. WO-4, 31) p.> ~KIMMEY, J. W.,' and

P. C. LIGHTLE. 1955. Fungi associated with cull in red-wood. For.
Sci. 1:104-110. LOWE, J. L. 1966. Polyporaceae of North America.

The genus Poria. State Univ. Coll. of Forestry, Syracuse Univ. Tech.
Bull’ 90, 183 p. NOBLES, M. K. 1948. Studies in forest pathology.
VI. Identification of cultures of wood-rotting fungi. Can. J. Res.,
Co, 26:281-431. NOBLES, M. K. 1958. Cultural characters as a guide
to the taxonomy and phylogeny of the Polyporaceae. Can. J. Bot.
36:883-926. NOBLES, M. K. 1965. Identification of cultures of wood-
inhabiting Hymenomycetes. Can. J. Bot. 43:1097-1139. NOBLES, M. K.,
R. MACRAE, and B. P. TOMLIN. 1957. Results of interfertility tests

on some species of Hymenomycetes. Can. J. Bot. 35:377-387. SINGH, B.
1966. Studies on Indian Poria. II. Diagnosis of five species as new
records. Indian Forester 92:680-683. STALPERS, J. A. 1978. Identi-
fication of wood-inhabiting Aphyllophorales in pure culture. Centraal-
bureau voor Schimmelcultures. Studies in Mycology: 16, 248 p.

MYCOTAXON

VGieecl Lint NO.e eee Dt eee or April-June 1981

A NEW SPECIES OF AMANITA
DAVID T. JENKINS

Department of Biology, University of Akabama in Biuningham,
BuUuntngham, AL 35294

ABSTRACT

Amanita martginata (fig. 1) is described as new from Tennessee.
It is assigned to Section Lepidella, Subsection Solttariae.

Amanita martginata Jenkins, sp. nov.
Holotype: Tennessee, Loop Road, Cades Cove, Great Smoky Mountains
National Park, 17. viii. 72, David T. Jenkins 580(DTJ).*

Pileus 110 mm mense diametro, convexus, margo leviter appendicu-
latus, non striatus; labrum distinctum sterileque subter, album conver-
tens ad gilvum colorem; reliquae volvae ita verrucae adnatae factaeque
in inaequalem formam, gilvae cum cacumenibus in vero colore, potius in
circlos qui easdem medias partes habent. Lamellae stipatissimae, sol-
utae. Stipes 90 x 20 mm, solidus, fibrillosus squameusque adversus
basem; tegumen ad apicem partim, pendens, album, delicatum; bulbus ad
basem in ovi formam factus, gilvus; maculans flammeus fuscusque adversus
fundum; aliquando rimae longitudine; reliquae volvae ita circli squam-
arum adnatarum reflexarumque leviter, qui easdem medias partes habent.
Sporae 5.46-6.25 x 7.0-10.15 um.

Fruit body medium to large, solitary. PILEUS: 110 mm diam, con-
vex, margin non-striate, very slightly appendiculate, but with a dis-
tinct, sterile margin, up to 3 mm wide, pileipellis fairly easily sep-
arable, fibrillose between volval remnants, white to creamy-white, flesh
white, up to 15 mm thick at center, tapering toward margin; universal
veil remnants as fibrous, adnate, irregular-shaped warts, up to 7 mm
wide at base, cream with tips cream to pale tannish-cream, more or less
concentric ring arrangement, becoming decidedly smaller toward margin.
LAMELLAE: very crowded, free, fairly broad, cream, edges smooth; lam-
ellulae numerous, attenuate. STIPE: 90 x 20 mm, solid, white, fibril-
lose-scaly beneath annulus, becoming lacerate-scaly toward base; partial
veil apical, creamy-white, pendant, large and thick, striate above,
floccose below, double-edged, very delicate, soon falling off, ring of
universal veil material approximately 10 mm below partial veil. Basal
bulb ovoid, cream at apex, staining orangish-brown toward bottom,

65 x 45 mm, occasionally with longitudinal splits; universal veil rem-
nants as concentric rings of adnate, slightly recurved scales, quite
thick, on upper part of bulb. Faint chloride of lime smell.

/PILEIPELLIS: filamentous hyphae densely interwoven, slightly
gelatinized, hyaline, 2-7 um diam. PILEUS TRAMA: filamentous hyphae

ar nn EES IEEEnEEeeed

* DTJ = The author's herbarium

ies

undifferentiated, moderately branched, clamps rare, 3-8 um diam;
inflated cells abundant, mostly terminal, variform, mostly elongate,

up to 162 x 31 um. LAMELLA TRAMA: bilateral; filamentous hyphae
undifferentiated, moderately branched, no clamps, 3-8 um diam; inflated
cells terminal, elongate. SUBHYMENIUM: hyphae ramose, clamps rare.
BASIDIA: up to 48 x 3.9-8.6 um, 4-sterigmate, thin walled, without
clamps. UNIVERSAL VEIL: filamentous hyphae on pileus moderately abun-
dant, sparsely to moderately branched, without clamps, 3-6 um diam,
irregularly disposed; inflated cells abundant, globose, subglobose,
broadly elliptic, elliptic, clavate, cylindric, most being quite small,
not exceeding 62 x 46 um, usually short, terminal chains, irregularly
disposed: tissue on basal bulb similar, but with a slightly higher
percentage of filamentous hyphae. STIPE TRAMA: filamentous hyphae
undifferentiated and abundant, sparsely branched, without clamps,

3-6 um diam; the presence of gloeoplerous hyphae conspicuous; inflated
cells terminal, clavate, longitudinally oriented, up to 312 x 25 um.
PARTIAL VEIL: upper surface mostly inflated cells, elliptic, ventri-
cose, clavate, up to 125 x 63 um, mostly terminal; filamentous hyphae
Sparse, sparsely branched, without clamps, 2-7 um diam: lower surface
Similar to that above, but with many more smaller inflated cells, vari-
form; filamentous hyphae also sparse, similar to above.

Fig. 1 Amanita marginata Jenkins 580(DTJ)

114

SPORES: 6 50460-6725. 7-0-10.15eum, (Ei lecorlmoc, eye Ba51)
elliptic to elongate, adaxially flattened, thin walled, hyaline, weakly
amyloid, spore print white; contents guttulate; apiculus sublateral,
short cylindric.

Habitat and distribution: terrestrial, mixed coniferous and
deciduous forest, Tennessee.

Collections examined: road cut, Loop Road, Cades Cove, Great
Smoky Mountains National Park, Tennessee, 17. viii. 72, David T. Jenkins
580(DTJ).

Discussion: Amanita marginata is assigned to Section Lepidella,
subsection Salitariae. This is due to the presence of such characters
as amyloid spores, an appendiculate pileus margin, a white to cream
pileus color, and variform volval elements.

As attempt has also been made to assign this organism to a part-
icular stirps. Following the organization of Bas (1969) this organism
would seem to belong to Stirps Strobiliformis, based upon the absence
of basidial clamps, white volval remnants with moderately abundant,
interwoven elements, and the production of irregularly shaped volval
warts on the pileus.

Although this organism is apparently related to the members of
Stirps Strobihiformts, only slight changes in a particular character
would allow it to fit into other stirps. For example, if the basidia
had clamps this organism could be placed into Stirps Vingineoides due
to its small to medium, detersile volval warts and the irregularly
disposed universal veil remnants, consisting of moderately abundant
hyphae and inflated cells. If the elements of the volval remnants on
the pileus had a more erect-parallel disposition, it would be more |
closely related to Stirps Polypynramis, due primarily to its color and
small to medium, detersile warts.

As the Amantta flora of the United States, and in particular the
southeastern region, is further studied the variation and diversity
becomes increasingly apparent. As can be seen with Amanita marginata
it is frequently quite difficult to determine the specific relationship
with other members of this section based upon the current classification
system. Therefore, the establishment of new stirps within this section
may be necessary in the future.

ETNERATURESCITED

BaSsmce 1969. Morphology and Subdivision of Amanita and a Monograph
on its Section Lepidella. Persoonia 5(4): 285-579.

MYCOTAXON

VO) eX ule NOM os EDD cg aLio nla Api June wLooL

CANDIDA PARATROPICALIS, A NEW SPECIES OF CANDIDA

JOHN G. BAKER*
Analytab Products, Inc.
Division of Ayernst Laboratonrtes
Plainview, New York 11803

TRAS ESS OALKIN
Division of Labonatonies and Research
New York State Department of Health
ALbany, New York 12201

DAVID H.) PINCUS
Analytab Products, Inc.
Diviston of Ayenst Labornatonrtes
Plainview, New York 11803

RICHARD F. D'AMATO
Diviston of Microbtology
Department of Pathology
Catholic Medicak Center of Braookkyn and Queens
Jamaica, New York 11432

An unusual Candida isolated from clinical specimens
has been found to be sufficiently different from existing
taxa within this genus to warrant its establishment as a
new species, C. paratnropicakrss.

MATERIALS AND METHODS

Twenty-nine isolates of the new Candida were recov-
ered from specimens of human blood, sputa, urine, bronchial
washings, throat swabs, a lung biopsy, and a decubitus
ulcer. Their morphologic and physiologic characters were
studied by procedures previously described (Baker, Salkin,
Pincus #eand DraAmato, Lo Silay 9D)

TAXONOMIC PART

Division: Fungi imperfecti.

Form class: Blastomycetes.
MOrMeOrdeL sm Crypetococcales.
RorvmEconihy, scl tOCOCCaceae.

Candida panatrzopreatis Baker, Salkin, Pincus et D'Amato,
St enOVem he LouUrem 1).

*Present address: Microbiology Department, Lahey
Clintc mpunl ngtonsMall, Burlington, Massachusetts, 01803:

Fig. 1. Candida paratropicakits on cornmeal + 1% Tween 80
agar vatters edaVseuncubati Onsate omc.

Coloniae in extracti malti agaro cultae 2 mm diam
usque sunt, ochroleucae farinulentae, lucidae vel subluci-
dae, planae subconvexae vel umbonatae, marginibus fila-
mentosis patulis. Per dies 14 in glucoso-fermenti extracto
cum aqua peptonica immersis sedimentum pelliculumque
formantur, cellulis fertilibus ovoideis vel elongato-
ovoideis (2.5--7.5 x 3.0--12.5 um) cum quibus blasticonidia
gemmantia numerosa collo angusto conjunguntur; pseudo-
hyphae hyphaeque genuinae adsunt. In farinae Zeae agaro
per dies 7 cultae fermenti cellulae ovoideae vel elongato-
ovoideae (2.5--6.5 x 3.5--14.0 um) cum pseudohyphis
hyphisque genuinis formantur. Status sexualis ignotus.
Notulis physiologicis (tab4 1)\ cum. (Ci 2r0opceals (Gast. )
Berkhout congruit, sed sucrosum melezitosum fermentare
necnon L-arabinosum assimilare nequit simulque methyl-D-

glucosidum sucrosum melezitosumque modo variabili assimi-
at.

Habitat: Man.

Holotype: 79MR8, isolated from human blood at autopsy,
New York, New York.

The holotype slide preparations, as well as living
cultures, have been deposited in the American Type Culture
Collection, Rockville, Maryland (accession number ATCC
42678), and in the New York Botanical Garden Culture
Collection, Bronx, New York.

The epithet C. parxatnopicakis was chosen to emphasize
the resemblance of this species to C. tropicalis.

Ley

Colonies on malt-extract agar at 14 days are 2 mm in
diameter; yellowish-white; pasty; glossy to dull glossy;
and umbonate, flat, or slightly convex, with spreading
filamentous margins. At 14 days in glucose -- yeast-
extract -- peptone water a sediment and pellicle have
formed; reproductive cells are ovoid to long-ovoid (2.5-
Pe Oe es OK. Omi) wi tCoemuLte ple budding, blastoconidia,
each attached by a narrow neck; and pseudohyphae and true
hyphae are present. When grown on cornmeal agar for 7
daysymeene celisvare ovoid! to Long-ovoid (2.5-6.5 x 93.5-
14.0 um) with well-ramified pseudohyphae and true hyphae.
The teleomorph is unknown. Physiologic characters (Table
1) are similar to those of C. tnopicalss (Cast.) Berkhout
excepLelom(1) the inability of C. paratrzop.calrcs to fer-
ment sucrose and melezitose and assimilate L-arabinose and
(ii) its variable assimilation of methyl-D-glucoside,
Sucrose, and melezitose.

DISCUSSION

The morphology of C. panratnopicakis (shape and size
of blastoconidia, development of blastoconidia in sparse
clusters at hyphal nodes, absence of chlamydospores and
germ tubes) suggests a close similarity to C. troptcalts
and its sSucrose-negative variant (Ahearn, Meyer, Mitchell,
NICHOLSON wednasloreanim~, 097 /-) baker etsal., §19:31a)) sHow-
ever, the new species' inability to ferment sucrose and
melezitose clearly differentiates it from C. troptcalrs
(van Uden and Buckley, 1970). The utilization of methyl-
D-glucoside, melezitose, and sorbose by many isolates of
C. panatnoprcaliss (24%, 52%, and 90% respectively; Baker
et al., 198la) and its inability to assimilate inulin
distinguish this taxon from the sucrose-negative form of
CEeLLODL CAC CA Anearn etials, 1977).

Since isolates of C. panatroprcakts frequently (41%)
do not assimilate sucrose, cursory examination of such
isolates could contribute to their misidentification as
C. 4tellatoradea.

The lack of recognition of this new species despite
its relative frequent recovery from diverse clinical
specimens in several laboratories suggests that C. panra-
tnopteakss has been and probably still is being misidenti-
fied.

ACKNOWLEDGMENTS

The authors express their appreciation to Dr. Rupert
Barneby for preparation of the Latin translation of the
HtaGnNOSIGNatOnDrSs Gi AvuandsandsGa eb. Roberts atom sup]
plying some of the cultures used in this study, and
especially to Dr. Michael R. McGinnis for providing taxo-
nomic assistance.

118

TABLE 1. FERMENTATION AND ASSIMILATION PATTERNS OF 29
C. paratropicalis ISOLATES?

Tsolates positive (3%)

Substrate Fermentation Assimilation
Cellobiose 0 69
Dextrose 100 100
Galactose 100 100
inwian 0 0
Lactose 0 0
Maltose 100 100
Melezitose 0 52
Melibiose 0 0
Raffinose ) 0
Sucrose 0 59
Trehalose 100 100
D-Arabinose 0
L-Arabinose 3)
Cierrceacia 100
Dulce 0
Brevcnienco | 0
Ethanol 100
Ethylamine

hydrochloride 100
Glucitol 100
Glycerol 14
THOS oll 0
2-Keto-gluconate 100
DL-Lactic acid 86
Mannitol 100
Methyl-D-glucoside 24
Potassium nitrate 0
Ribitol 100
Ribose 10
Rhamnose 0
Salicin 24
Sorbose 90
Succinic acid 100
Xylose 100

Results obtained with Wickerham broth technique at 3
weeks.

LITBERATURESECLTED

Ahearn, D. G., S. A. Meyer, G. Mitchell, M. A. Nicholson,
and A. I. Ibrahim. 1977. Sucrose-negative. variants
of Candadaltuopi. calc so. Clin Microbiol.

5: 494-496.

119

Baker MU lmG Wales! Soa ki nD tage LnCuUS and Reg ks:
D'Amato. 198la. Diagnostic characters of an atypi-
CamCanalid eC line Mi CroObLol. (in press).
Dakar Gomer moa Lk in De ieee 1 NGUS jj eand Re be
D'Amato. 198lb. Use of rapid auxanographic proced-
ures for recognition of an atypical Candida. J.
GloneeMLeroo.O bm ULE GOSS Jr.

van Uden, N., and H. Buckley. 1970. Genus 2. Candida
Berkhout, peo9 3206 /ee inn) seLoddersed.)7,) The
yeasts, a taxonomic study, 2nd ed. North-Holland
Publishing Co., Amsterdam.

MYCOTAXON

Wioiliee Xe NOt alee DOeel 20 bce5 April-June =voSL
ca. Eee ,
NOTES ON

CORTICIACEAES CBASIDIOMYCE Geapeav tke

Two new species of Tubulicrinis.
Kurt Hjortstam

Malaregatan 12
S-441 35 Alingsas, Sweden

SUMMARY

Tubulicrinis cinctoides and |l-povalisporus
are described from North Europe and Africa
respectively. Jhe first species is similar
Ome CUnCLUG eG sh CUM anu trae Ll ihinbe Cmy
its more globose spores and cystidia of di-
vergent appearance. The African species is
related to the I. glebulosus-—qroup in its
ecystidial morphology, but well defined py
having oval spores. The type of T. cinctus
was studied.

LUBULITCRINT SS CINCTOLIDES Hjortst. nO Vass eC usu Commer or.

Species Tubulicrini cincto affinis sed sporis oblique sub-
grobosis @4.75—)5 x0 (8.75-)4-4 5ovums el yocysGuci tom nds —
stinete capitatis, 60-/70(-90) x 5-6 um.

Holotypus: Sweden. Ostergdtland. V. Tollstad par., Omberg,
Storpissan state forest reserve, in herb-rich Picea forest,
on ‘decayed trunk of Picea abies. 1979-11-09. K Hjgortstami&
ie ad en gbac kt.weH jen clea ben Ge).

Paratypus: ‘do. Hjm 11393 (priv.herb.).

Rruitbody resupinateswetfuse, atetirstyreti culate mtLnen

more or less continuous, (thin, whitish «Wi che thescysttaia
projecting, each with a globule of excreted matter easily
seen under a lens (50-100 X).

Subiculum very sparse, composed of thin-walled, more or

less uniform hyphae, about 2-2.5 um wide, with a clamp at
each septum.

Lyocystidia faintly amyloid, mostly greyish in Melzer’s
reagent, cylindricai, up to 60-70(-90) um long and 5-6 um
wide near the bi-, seldom tri-furcate base; the capillary
lumen ending more or less abruptly; neck-width (3.5-)4-5 um,
the apices of the cystidia slightly expanded but indistinct-
ly capitate, usually not more than 6-7 Um*in diam.

Basidia small, 12(-15) x (4.5-) 5 (-6) um, with four sterig-
mata and a clamp at the base, not amyloid.

Spores obliquely subglobose, thin-walled, with neither amy-
loid non eyanophilous geaction, (4.75-) 5, x) (oe7>— ea 4

Atal

|
5 um
e.

ee |

5 um
-—
5 um

oO
09)

Fig.1. Tubulicrinis cinctoides a) cystidia b) basidium

¢) spores. = Coll. Hjm 11348 (holotypus). T. cinctus d) hy-
pha,e) ecystidium f) spotes.’— Coli. Cunn. 17428 (holotypus).

um, with a distinct apiculus.

Remarks. The new species T. cinctoides is undoubtedly very
Sina late Lomlomc nc tuseG.H.CUnn a bDubesSeemsmro beswel igide li —
mited by its more globose and slightly larger spores. In

the type of the latter (as far as known the only material
gathered) the spores are more or less ellipsoid and measure
WA Ani a1) Xe2 1 9-5 62503) UM (Cine thesoriginel. descrip=
tion 4-425 x 3-5.5-um), while most spores of 1. cinctoides
measure 4.75-5 x 3.75-4 um. The lyocystidia of the two spe-
cies show a slight but clear difference. In T. cinctus the
apical bulb is fairly distinct, starting somewhat ‘abruptly
from the neck (fig.1,e), and thick-walled in the basal part,
while the cystidial-bulb of the new species is less marked
and usually consists only of the thin-walled part of the
apex (fig.1,a).

122

7 —
Sym
sae
Ne b
ite
oN
j—-__|
C) 5 um
o Cc)
C
a

Fig.2. Tubulicrinis ovalisporus a) cystidia b) basidium
c) spores. - Coll. Ryv. 11308 (holotypus).
TUBUPTCRENTLOSOVAE FSRBURUSEHYOrtGte n0V .mSDe.c. mn 1 mecr.

specs ese TuUbULICrInIe Gglebulosomaltinis esedmiructincatLtone
subtiliter odontioide et sporis ovalibus 4.5 (-5) x 2.5-
2.75 (-3) ums; lyocystidiis cylindricis, pallide amyloidibus
(fere griseis), circiter 100 x 5-6 um, versus apicem levi-
ter angustum.

Holotypus: Africa. Malawi. Southern Prov., Mulanje distr.,
Mulanje Mts., Lichenmya Plateau, alt. 1800-2000 m.a.s.l.,

on coniferous wood, probably Widdingtonia. 1973-03-09/10.
L. Ryvarden 11308 (0). Isotypus: in GB.

Panatvypusi do wcll. Any Varden. o0 2mula.

123

Fruitbody resupinate, effuse, thin, composed of very small
aculei, about 15-20/mm, whitish or becoming yellowish brown,
with numerous cystidia projecting above the hymenium and
easily observed under a lens (50-100 X), each encrusted at
the apex with brownish, excreted material.

Subiculum thin, mostly inconspicuous, composed of thin-
walled, uniform hyphae with clamps, usually 2-3 um wide.
Lyocystidia numerous, cylindrical, faintly amyloid, about
100 um long and 5-6 um wide near the bi-furcate base, ta-
pering slight towards the apex and with a neck-width of

4-5 um, the capillary lumen ending gradually at the slight-
ByeLAape lung DULeOLULUSe, aplCal part.

Basidia usually 13-15 x 4.5-5 um, with four sterigmata and
with a basal clamp, not amyloid.

Spores ovate or ellipsoid, thin-walled, 4.5 (-5) x 2.5-2.75
(-3) um, with neither amyloid nor cyanophilous reaction.

Remarks. Owing to the cystidial morphology this new species
seems to be closely related to T. angustus (Rog.& Weres.)
Donk and T. glebulosus (Bres.) Donk but is distinguished by
MvuomOovatemuomellipsoi1dmespores. shes tinely sodontroid 7 ructi-—
fication is somewhat unusual in other species of the genus,
but recently Jilich described one species (T. corneri)

WO iCnem according tomthesdescription.aismquitesclosemto I:
hamatus (Jacks.) Donk but distinctly toothed, with fairly
Largeeaculei ma nmnuresnamaverta le mastmilaregrowth suructure
can be demonstrated for example in T. inornatus (Jacks. &
Rog.) Donk,where it is less pronounced than in T. ovalispo-
rus.

Acknowledgement

The author is grateful to Carl Stenholm’s foundation for
supporting the investigation of the Storpissan nature re-
serve. In a subsequent paper the region of Omberg will be
Breatedumore thoroughly.

References

Cunningham, G.H. 1963. - The Thelephoraceae of Australia and
NeWweeleal land. N./@.mUeD.) SCI ..51NGUSEI. RES DU LI.
145. pp. 142-143.

Donk, M.A. 1956. - Notes on resupinate Hymenomycetes - III.
Fungus 26: 3-24.

Jackson, H.S. 1948. = Studies of Canadian Thelephoraceae. |.
Some new species of Peniophora. Can. Journ. Res.
Ep Pa 45S) SeWateh Ue

Jilich, W. 1979. - Studies in resupinate Basidiomycetes - VI.
On some new taxa. Persoonia 10(3): 332-333.

MYCOTAXON

Vole DieweNO wel ep pee le 4126 Apral-June sy lost
BE ee ee eee
NOTES ON

CORTICIACEAE (BAS EDIOMYER TES aie

Three new combinations in Hypochniciellum.
Kurt Hjortstam

Malaregatan 12
S-441 35 Alingsas, Sweden

SUMMARY

The genus Hypochniciellum with the type spe-

cies Leptosporomyces ovoideus Jiilich was de-
scribed by Hjortstam and Ryvarden (1980) and
characterized by pellicular to membranaceous
fruitbodies and thick-walled, cyanophilous
spores.

Three new combinations are proposed here: H.
cremeoisabellinum, H. molle, and H. subilla-
queatum. The generic circumseription is
slightly emended to include also species with
spores greyish but not distinctly blue in Mel-
ZeOT Seereagen ti.

INTRODUCTION

When Eriksson and Ryvarden (1976) treated the genera Lepto-
sporomyces and Leucogyrophana they discussed several simi-

larities between Leptosporomyces ovoideus Julich and Leuco-
gyrophana cremeoisabellina (EitschsomPart.y. ame hbicmcenm
Parm., and L. subillagueata (Litsch.) Jiilich. The authors

also expressed) theypossibility of joinindgmechesspeci ese nea
separate genus.

Later on Ginns (1978) circumscribed the genus Leucogyropha-
na and excluded the three species mentioned above without
any suggestions as to their generic position. Consequently,
the species needa place in the family Corticiaceae, and in
my opinion Hypochniciellum seems to be suitable.

Without any doubt the thickness of the spore-wall is a cha-
racter to consider, uniting the species. In spite of the
fact that the spores of H. ovoideum are non-amyloid in com-
parision with the other species, which have a more or less

greyish reaction, the species have many characters in comm-
on

The genus Amyloathelia Hjortst. (1979) is in many respects
not appropriate for the species here referred to Hypochni-
ciellum. All species in that genus have strongly amyloid
spores and thinner spore-walls.

WAS)

Key to the species’ of Hypochniciellum

1. Cystidia present, basal hyphae with thickened

Vici isreme OO tae — OMI MEW. Cis tad. Gets lade teers te sei sede s «cs Soe OL Le
1. Cystidia absent, basal hyphae thin-walled
We Mey CSS) Ueto s Coho 6 Sh COD.S 6 tas DOI ae 2
POD OLE Sa Oli 7a UMenl OL Oiev.t-tsloretet<leve a «Js, c¥s1s fayecese H. cremeo-
isabellinum
PO DOT CSE — 45 58 mbar UMAl OIG cone ote ole oes tele sles sie. J
Spores UunchangedsineMelzerasereagent.. Known
Oa? ieieyoy Glebe Kelsiehs ood jaaavauoun cumeoc H. ovoideum
3. Spores greyish in Melzer’s reagent. Known
Oe Veet OM eC OL iis TOU SHOW OO Cisdststetets se ohelsielscey sess Sad lle yee
queatum

Pen CHUN Mele UMS OniS Dec mnyV sreenlyCOUCAXOM a2... 1s) 1) 6),0eh9.00 .
Type species: Leptosporomyces ovoideus Julich

Emendation.

Fruitbody resupinate, effuse, pellicular to membranaceous,
with a whitish, fairly well-developed subiculum, consisting
of loosely interwoven hyphae; hymenium smooth or with small
granules; margin inconspicuous or in some species fibrill-
SSeecvoOerhaZzomarphic.

Hyphal system monomitic. Basal hyphae thin-or more or less
thick-walled, subhymenial hyphae thin-walled, all hyphae
with clamps.

Cystidia present or absent, somewhat hypha-like, with thin-
Sreunvckened walls

spores thick-walled, cyanophilous, in some species with
Walls greyish in Melzer’s reagent.

BV OCHNLC TERE UM SGREMEOQISABELLINUMM@(Litsch.) Hjortst. nov.
comb. Basionym: Corticium eremeoisabellinum Litsch., Ann.
ic ON 2 = Sete 1 too

Lectotypus: Sweden. Sddermanland. Near Saltsjobaden, on Pi-
Rue sy Vestriss, 1906-11-14— LC. Romell (S). 2

this is a little-known species, but during recent years a
few collections have been found in Norway. One of these
specimens (Hjm 10530), differs in having spores greyish in
Melzer’s reagent and in having slightly encrusted hyphae.
It is possible that the species consists of more than one
baxom DULathis cannot at present be satisfactorily solved.

Selected specimens studied: Sweden. Smaland. Rumskulla par.,
Norra Kvill Nat. Park, on coniferous wood. 1770-10. Gro Gul-=
jenes.m.. (Ge and 0). Norway. Oppland. Dovre. Hjyerkinnholen,
elev. 950 m, on Pinus sylvestris. 1979-08-25. Hjortstam
oOo iil Ooo Om Cboth mune jMapbiv<therbee..

Be UC OND OLE GCUMeMOL ECE. aH jorts ti. noVes. comp.

Basionym: Thelephora mollis Fr., Syst. mycol. I, pag. 443,
1821.

Authentic™’material not located.

126

The modern interpretation of the species corresponds well
with the protoloque by Fries which reads as follows: effuse,

carnoso-membranacea, rubescenti-pallida, subtus tomentosa,
papillis prominulis majusculis.

On well-developed specimens the small granules (which were
observed by Fries) on the hymenium are easily distinguish-
able, at least eunder avlenss\o0)X),sandilogetherawi thetic
cystidia, spores, and wider basal hyphae with thickened
walls, delimit the species well from others of the genus.

Selected specimens studied: Sweden. Vastergotland. Bergste-
na par., Korpas, on decayed timber (coniferous wood). 1977-
08-07. Hjortstam 8141. Norway. Akershus. Nannestad. Tdmte
farm, on coniferous wood. 1978-09-27. Hermansen & Hjortstam
s.n. (both in Hjm priv. herb.).+Hedmark. Grue. Meldalen; on
Piceasabies. £19/74—-10=1 2 leaky Varden also 2mCUh).

HYPOCHNICITELLUM SUBILLAQUEATUM (Litsch=]) Hjgartst = novi.combr

Basionym: Corticium subillagueatum Litsch., Ann. mycol. 39
(Ee Taye ern

Lectotypus:* Sweden. Stockholm. ULidingo,, on Pinusysylvestris.
1910-05-29. L. Romell 2068 (S). !

For a good description and comprehensive discussion of the
Species see, Erikssonmand shy Vardenets7o.

Selected specimen studied: Sweden.Uppland. Arentuna par.,
Storvreta, on decayed coniferous fencing. 1927-05-08. Seth
Bunde VigiNOgeooenC Goer

References

Eriksson, J. & Ryvarden, L. 1976. The Corticiaceae of North
PUP Opes 2nV.0 lrmtem OS Lor

Pres; Eo. 1821oeovstemasmycologicume: mrund.

Ginns, J. 1978. Leucogyrophana (Aphyllophorales): identifi-
Cationpofespecies lan JGUCI mb Ole GGN6 sito =o

Hjortstam, K. & Ryvarden, L. 1979. Notes on Corticiaceae
(Basidiomycetes) V. Mycotaxon 10(1): 201-209.

-"- -"- 1980. Studies in Tropical Cort-
iciaceae (Basidiomycetes) II. Mycotaxon 12(1): 168-
184.

Julich, W. 1971. Monographie der Athelieae (Corticiaceae,
Basidiomycetes). Willdenowia Beiheft 7: 1-283.

MYCOTAXON

Vole X il mNOCMINEDD onl 27 1156 April-June 1981

CLADOSPORIUM BANTIANUM AND ITS
SYNONYM CZADOSPORIUM TRICHOIDES

Michael R. McGinnis” and Dante Borelli*

Department of Hospital Laboratories?, North Carolina
Memorial Hospital, University of North Carolina at
Chapel Hill, Chapel Hill, North Carolina 27514, and

Universidad Central de Venezuela“, Instituto de Medicina
Tropical, Ciudad Universitaria, Apartado 2109
Caracas, Venezuela

SUMMARY

Examination of the type material for Cladosportum
banttanum (Sacc.) Borelli and the living type culture of C.
trtchotdes Emmons, as well as a number of other living
cultures, has revealed that these two taxa are conspecific.
Based upon priority, C. banttanum is the correct name for
this taxon.

INTRODUCTION

During the past several years, there has been con-
siderable controversy regarding Cladosporium banttanum
(Saccardo) Borelli, 1960 and its relationship to C.
trichotdes Emmons, 1952. Borelli (4) studied the type
material for Torula banttana Saccardo, 1912, which was
isolated from a human case of cerebral phaeohyphomycosis
by Banti (1), and concluded that 7. banttana should be
transferred to the genus Cladosportum, for which, he
proposed the new combination C. banttanum. Borelli also
considered C. trichotdes, which was likewise isolated from
a case of human cerebral phaeohyphomycosis (3), to be a
synonym of C. banttanum. Emmons et al. (5) have disagreed
with Borelli. They consider C. trtchotdes to be a dis-
tinct species and Torula banttana a nomen confusum. This
study was undertaken to reevaluate these two taxa and to

128

resolve the controversy surrounding C. banttanum and C.
trtchotdes.

METHODS AND MATERIALS

Living cultures were studied on potato dextrose agar
(PDA) after 2 weeks incubation at 25°C in the dark. Meas-
urements were determined from camera lucida drawings of
slide culture preparations.

Herbarium specimens examined. Exsiccatum labelled
499, Torula banttana Sacc., Herbarium Mycol. Orto Botanico,
Padova. tally, ab.) Asesaccardo: §5 iphotomicrograpismone..
banttana prepared by P. A. Saccardo; DMD-38, slide culture
preparation grown on PDA, isolated from sawdust mulch heap
via spleen tissue of hamster, Williamsburg, VA, prepared by
D. Dixon in 1977, received as C. trtchotdes (see NCMH 1145
for living culture); DMD-39, slide culture preparation
grown on PDA, isolated from sawdust mulch heap via testes
of hamster, Williamsburg, VA, prepared by D. Dixon in 1977,
received as C. trichotdes (see NCMH 1148 for living culture).

Living cultures of C. banttanum studied. NCMH 111 =
CDC B-1937 = NIH 8595 = IP 509, from human brain abscess,
sent to NIH by G. Segretain, Institut Pasteur, Paris; NCMH
112 = CDC B-1938 = NIH 8580, from human brain abscess, T.
Collette, Sayre, PA.; NCMH 113 = CDC B-1940 = NIH 8579 =
ATCC 10958 = CBS 173.52, type culture, from human brain
abscess, C. Binford, Baltimore, MD; NCMH 114 = CDC B-1941
= NIH 8598, from soil, P. Klite et al., Panama; NCMH 115) =
CDC B-2003 = NIH 8504, from chronic abscess in human

abdominal wall, J. Pereira, Washington, DC; NCMH 116 = NCMH
121 = CDC B-2283 = CDC B-1897, from brain abscess in a
cat, S. Jang et al., Davis, CA; NCMH 117 = CDC A-980 = NIH

8590, from human brain abscess, J. Barnola Duxans, Caracas,
Venezuela; NCMH 121 = NCMH 116 = CDC B-1897, from brain
abscess in a cat, M. Rinaldi, Davis, CA; NCMH 122 = CDC B-
1898 = ATCC 22649, from human brain abscess, J. Bennett,
FL; NCMH 474 = NCMH 767 = CDC B-2525, from brain abscess in
a cat, Biberstein, Davis, CA; NCMH 767 = NCMH 474 = CDC B-
252). Lromebrain abscessminwaecat. SauangneuEalmaDavic.
CA; NCMH 1145 = DMD-38, from sawdust mulch heap, D. Dixon,
Williamsburg, VA; NCMH 1146 = DMD-41, from sawdust mulch
heap, D. Dixon, Williamsburg, VA; NCMH 1147 = DMD-58, from
stump of Juntperus virginiana, D. Dixon, Suffolk, VA; NCMH
1148 = DMD-39, from sawdust mulch heap, D. Dixon, Williams-

Table 1. Measurements of Cladosporium banttanum
conidia in herbarium specimens

Specimen Average size (um) Range (um)
Exsiccatum 4.6 X 8.4 Bei /. 50k 0-1 3.0
labelled 499
DMD-38 sea 2.2-3.9 X 5.7-9.2
DMD-39 SIL OOF is Jes —G 1X 5-1-1420

Table 2. Measurements of Cladosportum banttanun conidia
after 2 weeks on potato dextrose agar at 25°C

Isolate Average size (um) Range (um)
NCMH 111 faey Me Leet 1.8-3.2 X 4.5-10.0
NCMH 112 PERS PY OT OP 2.1-3.6 X 4.8-10.5
NCMH 113 Sab) 26 Tops} 2.3-4.0 X 4.7-11.0
NCMH 114 fay) DATTA 2.0-3.4 X 5.0-9.2
NCMH 115 Phat shoe € og fay 2.0-3.8 X 5.0-13.8
NCMH 116 Phalth 2 eT 23-5000 4 O-ae5
NCMH 117 ZR ke Dee 2.0-3.5 X 4.6-10.6
NCMH 121 SHAO] ee) 2.1-6.5 X 4.0-8.5
NCMH 122 ARIA Sys 2.1-3.3 X 4.0-8.0
NCMH 474 Maps TEP SRS 21-32 1X Oo
NCMH 767 Seiko dines 2.1-4.0 X 4.1-12.0
NCMH 1145 Pais D8 aiyate: Pe — 3 ok 490
NCMH 1146 SBOP os cas 2.3-3.7 X 3.7-9.0
NCMH 1147 PEAS Chey Joel Es Pye 119. Che aI ete yas
NCMH 1148 S51, 90th a 2.0-4.4 X 4.0-11.8
NCMH 1151 SAILS. Ce yes 2.5-4.1 X 3.8-9.0
NCMH 1152 PeORkEO. O 2 1—=3 2k ae EE
NCMH 1168 LAE RES SIRS 2 1-35 1X eo
NCMH 1181 Fest Pax Pe 2.13 eid 4. eos
NCMH 1182 Sie’ Sys 22S e kk oUF. 0
NCMH 1186 Sha74 Oe TO) 2.5-4.0 X 5.0-14.6

130

burg, VA; NCMH 1151 = ATCC 24928, from human brain abscess,
P. F. Jurgensen; NCMH 1152 = ATCC 28255 = Crichlow 853,
from human brain abscess, D. K. Crichlow; NCMH 1168 = CBS
328.65, strom a dog, Gs A. dev Vries; “NCMH 1S lg=soMaioo7,
from human brain abscess, M. Hironaga, Otsu, Japan; NCMH
1182 = SM 1522, from human cutaneous abscess, M. Hironaga,
Otsu, Japan; NCMH 1186 = FCM 7819, from human skin lesion,
S. Amma, North Kerala, India.

DISCUSSION

Measurements of the conidia preserved in the holotype
(Fig: 1)-for C.. banttanum.are 4.6 X 8.4.1m Gol 7G e@ 5. 0-
13.0 wm) (Table 1), which vary a little from the measure—
ments included in the original description (6) for the
species, and those measurements recorded by Saccardo in
his notes (Fig. 2). Saccardo described the conidia as
..."conidiis longe et sat persistenter catenulatis (in
quaque catenula 5-10), concoloribus, oblongo-ellipsoideis,
8-11 X 5, levibus, intus granulosis, summis junioribus et
pallidioribus.'' Even though Saccardo makes no statement
concerning whether or not the chains were simple or branched,
his photomicrograph of C. banttanum (Fig. 3) clearly shows
that the chains were branched. Figure 3 also reveals that
some of the chains consisted of more than ten conidia. The
exsiccatum contains a few blastoconidia that have three
hila, which confirms that the chains were branched.

Emmons et al. (5) have interpreted the original
description of C. banttanum as follows: "Saccardo stated
that 7. bantiana produced conidia.c) towLl 51 neu
branched chains which were only 5 to 10 cells in length".
Their translation of the Latin diagnosis for C. banttanum
is apparently in error since Saccardo considered only the
conidiophores to be simple. He described them as
...'conidiophoris ex mycelio hypostromatico brevi, cellu-
loso ascendentibus, fasciculatis, cylindraceis, simpli-
cibus, subrectis, variae longitudinis, plerumque 15-30u

Figs. 1-2. Cladosportum banttanum. 1. Lactophenol prep-
aration of exsiccatum labelled 499, Torula banttana
Sac., Herbarium Mycol. Orto Botanico, Padova, Italy,
P. A. Saccardo, in differential interference contrast

microscopy. 2. Notes prepared by P. A. Saccardo for
Torula banttana Sacc.

131

Py pO ay, OP ays

“rsppitane i To er

ith

eeSpagygage

OM tre

CO™
Yigg

152

long, raro usque ad 60y (ex cultura)..." Owing to their

incorrect translation of the nature of the conidial chains
for C. banttanum, they stated: "If the name, 7. banttana,
is based on both Saccardo's descriptions and Borelli's
interpretation of a photomicrograph attributed to Saccardo,
then the name, 7. banttana, apparently is based on two
fungi and is a omen confusum. Figure 3715 5thegpioco—
micrograph published by Borelli in 1960 (4) that is re-
ferred to by Emmons) e& al.) ltsclearlyssnowssthaeaenc
chains were branched and that only one fungus was involved.

It has been argued that C. banttanum and C. trichotdes
should be maintained as separate species because of dif-
ferences reported in the number of conidia composing the
chains (5). Emmons in Binford et al. (3) described the
conidia of C. tricnordes, ini part, as) ConidiisseMmipsoidis
vel cylindraceis, 2-2.5 X 4-10u, continuis vel l-septatis,
fuscis, aequalibus, formatis in cateniis longis ramosisque."
No mention was made by Emmons of the length of the conidial
chains other than that they were long, but his figure 5
shows the longest chain to posses six conidia. Examination
of NCMH 113 (Figs. 4-5), and additional cultures have
revealed that the chains are much longer, often consisting
of 35 or more conidia. Even though Saccardo stated that
the conidial chains of 7. banttana consisted of 5-10
conidia, his photomicrograph (Fig. 3) shows that some of
the chains were longer.

Emmons et al. (5) have observed that isolates ofuc,.
banttanum ..."have been remarkably uniform in morphology
and size of spores, although a few strains have borne
larger spores in primary cultures than those described by
Emmons, reverting to spores of typical dimensions upon
subculture.'' A comparison of DMD-38 and DMD-39 (Table 1)
with NCMH 1145 and NCMH 1148, respectively (Table 2),
Supports the observation that the conidia tend to become
smaller in size upon subculture. In a personal communica-
tion to Dr. Borelli, J. Bennett (National) Institutes of
Health, Bethesda, MD) studied a brain isolate that he

Figs. 3-5. Cladosportum banttanum. 3. Photomicrograph
prepared by P. A. Saccardo of Torula banttana showing
its microscopic morphology. 4-5. Branched chains of
blastoconidia (NCMH 113), potato dextrose agar after 2
weeks at 25°C, in differential interference contrast
microscopy.

15S

—

134

s+dentified as C. trichotdes. Bennett's isolate produced
conidia that were approximately the same size as those
described for C. banttanum by Saccardo. Unfortunately,
Bennett's isolate could not be included in this study
because it had died several years ago. After examining the
living cultures listed in Table 2, it is obvious that the
conidia of C. banttanum vary in size, ranging from 1.8-6.5
X 3.7-14.6um. The size range seen in the exsiccatum is
Bel -7 38x) > o-b3h01m.) which is compatiblewwithethervaria]
tion observed in the living isolates. This similarity in
conidial size clearly supports the conclusion that C.
banttanum and C. trtechotdes are conspecific, even though
the conidia in the exsiccatum tend to be larger than those
in the living cultures.

There is no doubt that Banti's isolate had the ability
tOmerowsat3/°Gu(0).n Inetact, Bantisroundstiataniomrungus
was pathogenic for rabbits (7), whose body temperature is
approximately 40°C. Since there is no living culture of
Banti's isolate, it is impossible to determine whether it
could grow at 42-43°C. Borelli (4) demonstrated that
isolates of C. banttanum can grow at this high temperature.
The thermotolerance and neurotropism in man of the original
isolates of C. banttanum and C. trtchotdes add additional
support to the conspecificity of these two taxa.

Another controversial point is the reported color of
the human brain lesions. According to Banti's description
(1), the cerebral nodules were ..."of a deep brown color,
giving one the impression of a melanotic sarcoma." Ac-
cording to Emmons et al. (5), the lesions caused by C.
trtchotdes occur as gray abscesses. Judging by what is
published in the literature, this aspect of the disease is
variable. For example, in a Venezuelan case (2), the
lesions were described as gray-blackish. In the necropsy
protocol of Banti's case, which was reproduced by Stigliani
(7), the lesions were described as a ..."dirty gray... from
which a thick, mucous thread-like, grayish-greenish liquid
came out...". The latter text was most likely dictated by
Banti during the performance of the necropsy. Since the
color of the lesions could have darkened within a few
hours, this may have resulted in the different descriptions
of the lesion color. The contention by Emmons et al. (5)
that the pathological aspects of the two infections are
Significantly different enough to maintain C. banttanum
and C. trichotdes as separate taxa is unjustifiable.

155

It is apparent that C. trichotdes and C. banttanum are
conspecific. The arguments put forth by Emmons et al. for
maintaining these as separate taxa are not compelling.
Cladosportum banttanum can be characterized as growing at
temperatures up to 42-43°C and by producing long, sparsely
branched chains of blastoconidia consisting of as many as
35 or more conidia. The conidia are smooth, 1l-celled
(rarely 2-celled), oval to oblong-ellipsoidal, 3.0 X 6.4um
(1.8-7.3 X 3.7-14.6um), and pale brown. The conidia arise
from hyphae or simple septate conidiophores that are pale
brown in color. The nomenclature is:

Cladosportum banttanum (Sacc.) Borelli, Riv. Anat. Patol.
OnGol ear Oo lOnmeLo OUR

=EOMU LG EDC LLAMA ECACC ANIM Y.COlem LO: O20 mL ole
(basionym)

= Cladosportum trtchotdes Emmons in Am. J. Clin. Pathol.
22:540-541, 1952.

ACKNOWLEDGEMENTS

The authors wish to thank Dr. Donald P. Rogers for
reviewing our manuscript, Professor Sergio Chiesa for loan
of the herbarium material maintained in the Saccardoan
Herbarium, and Dr. Dennis Dixon for loan of his slide
culture preparations and living cultures.

LITERATURE CITED

ame baiGinmG alo lle SOpLasuneCcasOndisO1td omicosiscere—
brale. Atti Accad. Med.-Fis. Fiorent. p. 49.

Mo yey se lLOLUS es Soprasun -caso7dis0idaomicos:
cerebrale. Lo Sperimentale 65:341.

ae Barletta.) wandeA.sAnCULOMOLLe Cameo lame GlLados
sporiosis profunda. Mycopathol. Mycol. Appl.
15:422-428.

3. Bintord Ge. so R.th-) lhompson, M. ha sGOtnameand. G-
W. Emmons. 1952. Mycotic brain abscess due to
Cladosporium trtchotdes, a new species. Am. J.
Clinweratiolse22:D30—542.

136

Borelli, D. 1960. Torula banttana, agente di un
granuloma cerebrale. Riv. Anat. Patol. Oncol.
yO loaO22.

EMMOUGS ComWeG emi DLULOLG me bem UCZ moni cme.
Kwon-Chung. 1977. Medical Mycology. 3rd. Ed.
Lea and Febiger, Philadelphia. pp. 471-482.

Saccardo, Py. A. 1912.) Notae mycologicae. lie fungi
ex Gallia, Abyssinia, Japonia, Mexico, Canada,
Amer. bor. et centr. "Anns Mycol .910:510—-3227

Stigliani, R. 1961. Particolarita istologiche della
torulosi encefalica nei preparati originali
Bantiani della prima Osservazione conosciuta.
Arch. Vecchi, Firenze 36:329-343.

MYCOTAXON

Foleex Ube aNowl le ppenl 572144 April-June 1981

A PRELIMINARY DISCOMYCETE FLORA OF MACARONESIA:
PART 2, HYALOSCYPHACEAE SUBF. ARACHNOPEZ1ZOIDEAE*

RICHARD P. KORF

Plant Pathology Herbarium, Cornell University
benacass Newmy OFkw1 GS 538USA

"Much like a subtle spider which doth sit
In the middle of her web, which spreadeth wide."
Sir John Davies
THE IMMORTALITY OF THE SOUL

Order HELOTIALES
suborder HYMENOSCYPHINEAE
Family HYALOSCYPHACEAE Nannf. 1932
Subfamily ARACHNOPEZIZOIDEAE Korf 1978
Tribe Arachnopezizeae Nannf. in Korf emend. Korf 1978

ONE KNOWN MACARONESIAN GENUS

ARACHNOPEZIZA Fuckel 1870 emend. Korf 1952
Key to the known Macaronesian species

1. Ascospores 5-7-septate at maturity. 3. A. aurata
1'. Ascospores O-1-3-septate at maturity ...--..seseee- 2

2(1'). Ascospores O-1(-2)-septate at maturity;
apothecia on husks of chestnut (Casta-
gteks))) “Welbbaety Gkoo gs oongupocdon.: 6 Dou oGo ue 3
2'(1'). Ascospores mostly 3-septate at maturity;
apothecia on wood, leaves, duff, acorns

3(2). Hairs thin-walled, tapering, usually 1l-septate,

* This flora will appear in an irregular order, with Part
1, containing the introductory material, one of the last to
be published. Reprints of individual parts will not be a-
vailable. References will appear in the final part. For a
geographical definition of Macaronesia and other special
features of this flora, see the note on page 144.

138

not constricted at the septum.
1. A. aranea f. aranea
3'(2). Hairs thicker-walled, multiseptate, constricted
Lnemseptar 2. A. aranea f. monilipila

4(2'). Apothecia 1-3 mm _ diam., ascospores
1LOR2=—135 2(= 162 le Kae? 2-3 amie

4. A. aurelia

ii( 2" j)2esApothecia, less, than O.5emmedian sare

5(4'). Apical cells of hairs without cyanophilic con-
tents; ascospores (13.5-)18.3-22(-26) x 2.9-3.7
ym; paraphysis apices variously deformed.

5. A. obtusipila

5'(4'). Apical cells of hairs with cyanophilic contents;
ascospores (14.6-)16.1-25.7 x 2.2-4.4 ym; para-
physis apices filiform, not deformed.

6. A. zonulata

1. Arachnopeziza aranea (De Not.) Boud., Icones Mycol.
Expl. Pl., sér. 3: 2. 1906 forma aranea.

= Lachnum aranea (De Not.) Lindau, in Engler
& Prantls Nat. Pflanzenfam: 11 (130)-
ADB. kelley

Arachnoscypha aranea (De Not.) Boud. ex Den-
nis weMycolsg Ra pe o288o me lo49..

RECENT TAXONOMIC TREATMENTS: Dennis (1949, 1978),
Korf (1952).

PREVIOUS MACARONESIAN RECORDS: None. SA,

TYPE LOCALITY: Italy. Qa |

KNOWN MACARONESIAN DISTRIBUTION

inks be reo A ae eee
Madeira. CUP-MM 1542. ’ ’
CANARY ISLANDS. yeni 1542 a ua

La Palma. CUP-MM 870.
SUBSTRATA: On cupules (burrs) of Castanea _ sativa.

Notes: I saw one 2-septate ascospore in my mounts (il-
lustrated), but typically the spores are either continu-
OUSS.Or sl-septate- Probably common wherever chestnut
grows, but difficult to find among the hairs on the in-

eressUlil aces ol (hemmspinyeenusks. wal cannot jagree with
Dennis, who separated this species in a distinct genus,
Arachnoscypha Boud., of which it is the type species,
but which Boudier himself later abandoned. In my earli-
er monograph (Korf, 1952), I missed Lindau's transfer
to Lachnum, which should be added to the synonymy
eivienmthere.

2. Arachnopeziza aranea (De Not.) Boud. f. MONILIPILA
KOGieaeieeEnhOVc

PREVIOUS MACARONESIAN RECORDS: None.

A Arachnopeziza aranea
f. aranea pilis multisep-
tatis miniliformibus dif-

Gent. 0
Differing from the type cular Se Ee

form in having multi- (}

septate rather than 1- TM Lene
SeptateCmelallS,mrands =bDy
the constrictions at the
septa, giving the moni—
TiO Gime S DCC Umma tLOmmaLhe

hairs. ae es

A. aranea f. moni-
lipila, ascus, pa-
raphysis apex,
7 spores, subi-

Oe CliG ah mn OU mesh mer OO Cl aml) mellelile Delta mm aMc
Kohn, on cupules of Castanea sativa, Castanea grove
below Pousada Vinhaticos, Madeira, Portugal, Wie Oe

KNOWN MACARONESIAN DISTRIBUTION
MADERIA.
Madeira. CUP-MM 1545 (holotype).

SUBSTRATA: On spiny burrs of Castanea sativa.
Notes: The asci arise from repeating croziers, and the
éctal excipular cells are quite large. The new form

may represent merely a growth-stage of the species.

3. Arachnopeziza aurata Fuckel, Jahrb. Nassauischen
Vereins Naturk. 23-24: 304. 1870.

RECENT TAXONOMIC TREATMENTS: Dennis (1949, 1978),
Nhat. (AESIEVAIh

140

PREVIOUS MACARONESIAN RECORDS:
*Dennis) corals (19/7).

iY Pe BOCAS TYee Germany

KNOWN MACARONESIAN DISTRIBUTION

LV oy Dai ie Vic
CANARY ISLANDS.

Gomera. CUP-MM 1348, 1368(TFC).

Hierro. CUP-MM 1446(TFC), 1450.

La Palma. CUP-MM 898.

Tenerife. CUP-MM 223, 230, 246,
256 ec0C CUE), eacD Ube )e
Sele mC, SykAg Wasrl

* AZORES.
*Terceira. “Dennis & al., Ws

SUBSTRATA: (On pwooduand: thunk:of Briccl nates manent

arborea, on bark of Myrica faya, wood Ao GS2 Dele
; SPS FTO : CUP-MM 1145,
of Castanea sativa, and undetermined Saisie

decorticated wood and bark.

Notes. This is surely the most common species of the
genus in Macaronesia, as it tends also to be in North
Americar andein ws burope.

4. Arachnopeziza aurelia (Pers. : Fr.) Fuckel, Jahrb.
Nassauischen Vereins Naturk. 23-24: 303. 1870.

RECENT TAXONOMIC TREATMENTS: Dennis (1949, 1978),
Korf (1952).

PREVIOUS MACARONESIAN RECORDS:
None.
iY Bre EOGAL IY =r cance. H
KNOWN MACARONESIAN |
DESERT BUss LON
CANARY ISLANDS. A. aurelia, 7 as-
Hierro. CUP-MM 1459 (OSC, cospores, CUP-MM
eG F459.) =x 1000;

SUBSTRATA: On fruits and husks of Castanea sativa and
other vegetable debris.

Notes: In North America and Europe this is most common
on cupules and leaves of Quercus.

141

5. Arachnopeziza obtusipila Grelet, Amateur Champignons
Olt) see corme oO eneemende Kot aMycologiaw43:@213%
1951.

RECENT TAXONOMIC TREATMENTS:
oul O51 O52).

PREVIOUS MACARONESIAN RECORDS:
*Baagge & al. (1972) as "inopercu-
ate discomycet..

fer LOCALITY = Erance.

KNOWN MACARONESIAN
DISTRIBUTION

*MADEIRA.

*Madeira. *CUP-MM 2502(C).

SUBSTRATA: On _ decorticated wood
of Pinus pinaster.

Notes aul ame inagebted™ tothe stu—
dents of the University of Copen-
hagen who collected this (Baagge
baal. ,o/2)8 ands for calling “the
Specimenmto my attention for iden—
tirication.

A. obtusipila, as-
cus, paraphysis, 5
ascospores, hate.
CUP-MM 2502,
xeeLOOO:

6. Arachnopeziza zonulata (Rolland) Boud., Hist. classi-
He, Ghicteleynyen Wiley jig We. ISON

RECENT TAXONOMIC TREATMENTS: Korf (1952).
PREVIOUS MACARONESIAN RECORDS: None.
TY PEMLOGCALITY: Corsica (France).
KNOWN MACARONESIAN DISTRIBUTION

CANARY ISLANDS.

Tenerife. CUP—-MM 1222.
SUBSTRATA: On wood of Pinus canariensis.
Notes: In the absence of type material, I (Korf, 1952)

Btreated this as a doubtful species. If I am correct in
interpreting this as Rolland's species, it is only criti-

142

cally distinct from A. obtusipila. Malencon and Bertault
(1958) proposed varietal status for Grelet's species un-
der A. zonulata, but their combination was not validly
published. I have made 3 trips to Rolland's type locali-
ty near Corte, in Corsica, but did not find any species
matching his description.

A. zonulata, section through the apothecial margin with
2 hair bases, 4 ascospores, paraphysis apex, ascus with
J+ pore, 3 hairs in cotton-blue, CUP-MM 1222, x 1000.

143

Tribe Polydesmieae Korf 1978
ONE KNOWN MACARONESIAN GENUS

Pol DEOMIAREBOUdTCH am BULlA es soOC. se Mycol. = Prance™ 1- 9113.
1885.

Key to the known Macaronesian species

Pm ScOSpores al—3—septate sat mmaturity,.9.0-13.5 x 2.2—

Be dh pola 1. P. fructicola
1®. Ascospores 3-septate at maturity, (13.9-)14.6-20.5
(-22.0) x 3.7-5.1 pm. 2. P. pruinosa

1. Polydesmia fructicola Korf, Mycotaxon 7: 475. 1978.

RECENT TAXONOMIC
TREATMENTS Kort
CUSH Ee

PREVIOUS MACARONESI-
AN RECORDS: “Konrf
(1978), +tBeltrdn Tejera
(1980).

ree LOCAL LT.Y: Ma-
deira.

KNOWN MACARONESIAN
DISTRIBUTION
*MADEIRA.

*Madeira. *CUP- Uf
MM 1504 (holotype) 7
(TFC), *1533, *1551, Vv
*1556, *1618, *1619,
#2333, *2361, *2362, — Dini San
2200 ea 2c9 |

*CANARY ISLANDS.
*Tenerife. *CUP-MM ae

1306.

SUBSTRATA: On pedun- P. fructicola, ascus with J+
cles, pods, twigs and pore, ascus with spores, para-
wood of Acacia spp., physes, 4 upper ascospores,
bark and capsules of CUP-MM 1504; 4 lower spores
Eucalyptus spp., un- CUP-MM 2361; all x 1000.
mdetermined bark . and

branchlet.

144

2. Polydesmia pruinosa (Jerd. in Berk. & BreveeBoudien,
Bull. Soc. Mycol. France 1: 113. 1885.

RECENT TAXONOMIC TREATMENTS:
Dennis (1978), Korf (1978).

PREV LOUS MACARONESIAN RE-
CORDS: *Korf (1978), t+tBeltran
Tejera (1980).

KNOWN MACARONESIAN
DISTRIBUTION

*MADEIRA.

*Madeira. *CUP-MM 1481(TFC),

1627) ay (OL ee LOO
*CANARY ISLANDS.

*La Palma. *CUP-MM 639

(le Ga ee / OO.

SUBSTRATA: 2On = pyrenomycetes,
on pyrenomycetes immersed in
wood, on branch of Acacia sp.

SPECIAL FEATURES OF THIS FLORA

Since, Party 1; thes introductory »material, 1s
to appear later, one reviewer of this part
confessed he did not know where Macarone-
sia is, and that he could not find it in
any of the atlases he consulted! Thus:
Macaronesia: the Atlantic island chain
comprising the archipelagos of the Azores,
Madeira, Salvage Islands, Canary Islands,
and Cape Verde Islands. Etymology: from

Greek, uakapos (fortunate) + vfigcos (is-

land): "Les Tfles Fortunes," 'The Fortunate .

Islands."" (Want to guess how often these P. pruinosa, 4 eee
papers will be cited as Macronesia instead spores, paraphysis
of Macaronesia?) apex, CUP-MM 1481,
Substrates follow O. Eriksson, A. Hansen SC LOO.

& P. Sunding. 1974. Flora of Macaronesia.
Check-list of vascular plants, 1974. Uni-
versity of Umea, Sweden. iv. + 66 pp.

Distributional symbols: Papers with previous Macaronesian records citing spe-
cimen data are indicated by one or more asterisks (* ** ***) and the cor-
responding specimens examined have been coded to those asterisks with their
CUP-MM numbers; where such specimens cannot be located or were not avail-
able on loan, the symbol ''n.v." follows brief specimen data. Papers citing
taxa without giving specimen data, or merely repeating previous records,
are indicated by a dagger or daggers (t+ tt ttt).

Herbarium abbreviations are those of the Index Herbariorum, ed. 6, and up-
dates appearing periodically in the journal Taxon.

Without the generous financial assistance of the National Science Foundation
(Grant DEB75-23557) this preliminary flora would never have been undertaken.

MYCOTAXON

WOR. XLT GNow Ll, pp. 145=149 April-June 1981

A PRELIMINARY DISCOMYCETE FLORA OF MACARONESIA:
PART 3, HYALOSCYPHACEAE SUBF. TRICHOSCYPHELLOIDEAE*

LINDA M. KOHN

Department of Botany, Clemson University
GlemsonwaeoouthsGaro linay79631,.USA

and

Plant Pathology Herbarium, Cornell University
Tthacas. New oYork 14853 USA

"A Hair perhaps divides the False and True."
Omar Khayyam (tr. Edward Fitzgerald]
RUBAIYAT, Stanza 19

Order HELOTIALES
Suborder HYMENOSCYPHINEAE
Family HYALOSCYPHACEAE Nannf. 1932
Subfamily TRICHOSCYPHELLOIDEAE Nannf. 1932
ONE KNOWN MACARONESIAN GENUS
LACHNELLULA Karsten 1884 emend. Dennis 1962

Key to the known Macaronesian species

1. Apothecia white, with a bright orange hymenium, on
living trees and cut ends of stumps of Pittospo-

rum spp. 1. L. pittospori subsp. azorica
ier Apothecia ~ogreenish-glaucus ' or grey ‘to’ nearly
BIS gononéocodsocaeto pd Oans Ao Sabon Coan oon anh fs

2(1'). Apothecia grey to black. 2. L. pulveracea
2'(1'). Apothecia greenish-glaucus, strongly pli-
Sater 3. L. viridi-glauca

* The parts of this flora, under the editorship of Richard
RB. Korf, will appear in irregular order. Reprints of indivi-
dual parts will not be available for distribution.

146

iz Lachnellula pittospori Kohn, Mycotaxon 12: 278.
1980, subsp. AZORICA Kohn, subsp. nov.

RECENT TAXONOMIC TREATMENTS: Kohn (1980). \

PREVIOUS MACARONESIAN RECORDS: None.

Cum subspecie typica Lachnellu- DIN \\

lae pittospori omnio conveni-
ens sed pilos erectos apice tu-
midos (in subspecie typica non
praesentes) ad marginem prae-
bens. Hi pili ex cellula apica-

et cellulam alteram supra eam
primam formantem. Huius subspe-
ciei ;novae ,distributio jam con—
nita ad Insulas Azoricas limi-
tata est. Holotypus: CUP-MM
2140.

li saepe pariunt processum fi- \
liformem aliquando tumescentum y

SS S55

L. pittospori subsp. azo-
rica, 5 smMarcinal hairs,
3 ascospores, CUP-MM
DIAZ. xe LOOGR

Agreeing with the _ descrip-—
tion of L. pittospori subsp.
pittospori in all respects ex-
Cepl, Lon ney presence solerc—
rect, “hairs with iswollen vapices vatwihewmaroinwenotspLe—
sent 1m) suUDSp. #=pltlLospori.s | bnese® ahains se oLvenmaproctce
from the apical. cell a filiform process, which in some
cases “swells to form “a second swollen sicell vabovemsthe
first. The distribution of the new subspecies is so far
as known limited to the Azores Islands. The type sub-
Species was described from Bermuda.

HOBO. Bem geal short mals cM meh Oils @ NoPE KOLO men ny mR OS ss
Man aeOle. ctUMDuL Ot meh tTLOSDOGUINGmS Dt mash | Det ha mm mene ill 2m
Lombas north of Caveira, Flores, Azores, ROctug a lye tos
iv.1978. (CUP-MM 2140.)

KNOWN MACARONESIAN DISTRIBUTION
AZORES:

Flores. CUP-MM 2138 (TFC), 2140 (holotype).
Terceira. CUP-MM 2009 (TFC).

SUBSTRATA: On Pittosporum spp., (stumps, cut areas of
living trees, and rotted wood of standing trees).

147

2. Lachnellula pulveracea (Alb. & Schw. : Fr.) Den-
nis perersooniar 222154281962"

RECENT TAXONOMIC TREATMENTS: Dennis (1949), Héhnel
M917).

PREVIOUS MACARONESIAN RECORDS: *Dennis & al. (1977).
ive eLOCALIDY: Germany.

KNOWN MACARONESIAN DISTRIBUTION

BA ZORES:

*Terceira. *CUP-MM 1681(K), 2011, 2382, *Dennis

Sra ees 1 Viel Sane

MADEIRA.

Madeira. CUP-MM 1560, 1621, 1623.
CANARY ISLANDS.

Gomera. CUP-MM 1343, 1354(TFC).

Hierro. CUP-MM 1424, 1444(TFC), 1477.

La Palma. CUP-MM 825.

Tenerife. CUP—MM 87, 248, 538, 1212(TEC).

SUBOERATA: On twig. of Aca—

cia sp., of Ilex perado sub-

Sp. platyphylla, of Ulex eu- (i
ropeus and Ulex sp., on
Stemivor Rubus®’sp-, ~and “on
eractermined "twigs, “bark,
wood, cut stumps, branches
and roots.

Notes: This distinctive spe-
cies, collected on various
woody substrates, is char-

acterized by apothecia that ?
turn violet in KOH and red- aly
piche ineiodine, was noted [N\Q

by Dennis (1949) and by
Héhnel (1917: No. 1020). As |, pulveracea, ascus,

mohnelweraiso: noted, “often nvei hair in KOH-
; paraphysis,

the asci are immature. The phloxine-glycerine, hair

apothecial structure is typi- in lactic-blue (CUP-MM

cally Trichoscyphelloideus, 87); 5 upper spores

mitnieed completely. celatini— (CUP-MM 1477); 5 lower

zea’ long—celled ‘éctal excipu-— spores (CUP—MM- 1354);
lum, unlike that in Hyalo- Allee LOO

= scyphoideae. In lactic acid-

148

cotton-blue the granulations on the hairs are no longer
visible. Raitviir (1970) treated it as a doubtful or ex-
cluded species of the genus Lachnellula, and Dharne
(1965) did not recognize it in the genus. Rehm's Ascomy-
ceten No. 1580, issued as Dasyscypha coerulescens var.
dealbata Rehm, was examined and found to be conspeci-
fic, confirming Dennis's (1949) synonymy.

3. Lachnellula VIRIDI-GLAUCA Kohn, sp. nov.

PREVIOUS MACARONESIAN RECORDS: None.

Apothecia gregaria vel fasci-
culata, viridi-glauca, sessili-
a, piulis -albissvestita se protun—
de cupulata, receptaculi parie-
tibus penitus plicatis praeser-
tim in apotheciis immaturis; re-
ceptaculum O.25-0.50 mm in
Gram.) scOommutatio. xanthochrouca
in 2% KOH eveniens. Excipulum
medullare “ex textura intricata
hyalina formatum, ss ce lu visee2—3

L. viridi-glauca, hair,
ascus and paraphysis,

me latis. WExcipudum = ectalege ex @
textura oblita formantum hyali- CUP-MM 850, x 1000.

na intertexta in gelatina copiosa flava contenta, cellulis
3-6 pm latis, exterioribus pilos edentibus. Pili hyalini,
erecti, septati,-grosse granulosi, 35=45 x 2 pm.,Asci elava—
ti, 36-45 x 5-6 pm , ex uncis iteratis enati, 8-spori, api-
ce magnopere incrassato, pori canalis pariete J-— cum/sine
usu KOH antecedente. Ascospori ellipsoidei vel interdum al-
lantoidei, 2-multi-guttulati, 6-7 x 2-2.5 pum. Paraphyses
filiformes, septatae, ramosae, ascos longitudine aequantes,
1 pm latae, in gelatina contente. Holotypus: CUP-—MM 850.

Apothecia gregarious to fasciculate, greenish-glaucus,
sessile, clothed in white hairs, deeply cupulate, walls
of receptacle deeply plicate, especially in young apothe-
cla; receptacle 0.25-0.5 mm diam;.xanthochroic reaction
in 2% KOH. Medullary excipulum of hyaline textura intri-
Cata, 7 cells™ 2-35 jim wide. jiictal excipuliummof yhiva liner
interwoven textura oblita bound in a copious, yellowish
gel, cells 3-6 pm wide, outer cells giving rise to hairs.
Hairs hyaline, erect, septate, coarsely granulate, 35-
fagy Re. PR MWTE Asci clavate, arising from repeating cro-
ziers, 8-spored, apex greatly thickened, pore channel
wall J- with or without KOH pretreatment, 36-45 x 5-6
pm. Ascospores ellipsoid or,occasionally allantoid, 2-

149

multiguttulate, 6-7 x 2.0-2.5 wm. Paraphyses filiform,
Bep.dleyem branched, = samen iength “as asci, 1 yam wide,
pound= ity ‘gel:

ool te meivete uh Oll, aN. GaebDenison, UsM. Kohn. & M.A.
Sherwood, on wood of ?Castanea sativa Mill., near mine
entrance at Km mark 13, road between Buenavista and
Piel asco,eeLassralma,. Canary —lslands, Spain,” 18.1.1976.
(CUP-—MM 850.)

KNOWN MACARONESIAN DISTRIBUTION
CANARY ISLANDS.
La Palma. CUP-MM 850 [holotype] (TFC, isotype).

SUBSTRATA: On wood (of ?Castanea sativa).

Notes: The greenish-glaucus color, the deeply plicate
receptacle (giving young apothecia the appearance of
quae bundtmcake!). andy the xanthochroice reaction in 2%
KOH makes this species especially distinctive. Neverthe-
less, the ectal excipulum composed of textura oblita
and the granulate hairs show Trichoscyphelloideus affi-
mities.) Note’ that young “apothecia often have no asci
and bear a copious gel layer above the hymenium.

MYCOTAXON

Volar Xt SaNoO wee pp alo 1.0 April-June, 1981

RESINOMYCENA GEN. NOV. (AGARICALES), AN ALLY
OF HYDROPUS, MYCENA AND BAEOSPORA

S.A. REDHEAD

Btosystemattes Research Instttute, Research Branch,
Agrtculture Canada, Wn. Saunders Bldg., C.E.F.,
Ottawa, Ontario, Canada KI1A 0C6

&
R. SINGER

Field Museum of Natural Htstory, Roosevelt Road at
Lake Shore Drtve, Chicago, Illinois, U.S.A., 60605

ABSTRACT

A new genus, Restnomycena, is proposed to accommodate
Agarteus rhododendrt, the tyve, Mycena kalalochensts, and
three new species, R. brunnescens, R. montana and
R. acadtensis. At present the genus is known with
certainty only from North America. The relationships to
the genera Baeospora, Hydropus and Mycena are discussed.

Agaricus rhododendrt Peck and Mycena kalalochensts A.H. Smith
along with three previously undescribed taxa form a clearly defined
taxon characterized by amyloid spores, pseudoamyloid tramal tissues
and a turf-like pileal epicutis composed mainly of resin secreting
oleocystidia.

In her monograph of the northeastern Marasmius species, Gilliam
(1976), excluded M. resinosus Peck (= A. rhododendrt) and suggested
that a new genus might need to be proposed for it. A.H. Smith (1947)
excluded both Omphalta rhododendrit (Peck) Sacc. and M. resinosus from
Mycena although at the same time describing M. kalalochensis as a new
Species of Mycena. These epithets were not dealt with in The
Agaricales in Modern Taxonomy, (Singer 1975). In this last
publication the species described here clearly key out to the
Tricholomataceae trtbus Myceneae and less clearly to the choice
between Hydropus and Mycena. However, the delimitations of both
genera exclude the group. Thus the recognition of a distinct genus
is a viable option. Alternatively emendation of Hydropus or Mycena

from the concepts accepted in The Agaricales tn Modern Taxonomy

could be proposed. The emendation of Baeospora by the inclusion of
Baeospora pallida Singer (1977), a species with close but not crowded
lamellae and white carpophores, is cause for considering this last
genus aS a possible depository for A. rhododendri and allies. As
detailed at the end of this paper, the erection of a new genus most
clearly resolves the problems surrounding the placement of these
species. Emendation of existing genera would make them too
heterogeneous.

Restnomycena Redhead & Singer, gen. nov.

Habitus omphaltodeus vel marasmiotdeus, superficetbus + restnosts
ex oleocystidits (frequenter tntermixtts elementis filamentosts vel
dendrotdets). Lamellis adnati vel subdecurrenttbus, confertis vel
subdistanttbus (haud confertissimis), ad actem oleocysttdits numerosts
tnstructis. Stipe aequalts, subearttlagineo, steco subcorneo, +
restnaceo. Tramate debtltter vel sat mantfeste pseudoamylotdeo
(parttbus subgelatinascenttbus exceptts); tramate hymenophoralt
regulart. Sports hyalints, amylotdets, levtbus, acyanophilts.

Typus: Agartcus rhododendrt Peck.

Basidiomes omphaloid or marasmioid, resinous, covered with
Oleocystidia intermixed with dendroid hyphae, white or dully pigmented.
Lamellae adnate to slightly decurrent, close to subdistant but not
crowded, with resinous edges formed by numerous oleocystidia. Stipe
equal, cartilaginous but drying to a horny consistency, resinous,
covered by scattered or clumped oleocystidia. Tramal tissues weakly
to strongly pseudoamyloid where walls not slightly gelatinized, but
frequently some inamyloid hyphae intermixed. Hymenophoral (lamellar)
trama regular. Spores smooth, amyloid, hyaline, acyanophilic.

Type species: Agaricus rhododendrt Peck.

A key to Restnomycena species

Pee ELUMPEULODS balce cisco cee e's eatatéle che eee sets cients ets .- Marasmtus rhododendrt

(see discussion)
A. From eastern North America ..... a iale siete tele maven 6 conte 5 ib retois esse eeia oe AE
Aeron western North America” ....c+ccce es a teroters ates es etatare ¢ Bienes ae ticle D

B. Pileus brown; hyphae incrusted with

B. Pileus white or whitish; hyphae lacking

Pigmented Aincrustation ....ccccccccesovecs Braletere afst tiene aie eis e aie a c
C. Lamellae close; spores small,
He An. 5 Ko Ano. (455) 1M. «ole oe RA nee CEO . R. rhododendrt

C. Lamellae moderately spaced to subdistant;
spores large, 9.4-12.8 X 4-5um ......---- eee Ee ITE Ph acddterners

u5z

D. Pileocystidia mainly capitate, secondarily septate,

mostly erect; on angiosperm litter ..........6.. R. kalalochensts
D. Pileocystidia clavate but rarely capitate, rarely

if at all secondarily septate and often collapsed

to a repent condition; on coniferous litter .......... R. montana

Restnomycena rhododendri (Peck) Redhead & Singer, comb. nov.
Figs. 1-5.
Agartcus rhododendri Peck (1875: 94)
Omphaltta rhododendrt (Peck) Saccardo (1887: 335)
Omphalopsts rhododendri (Peck) Murrill (1916: 311)
Marasmius decurrens Peck (1872: 77) nom. illeg. non. M. decurrens
Montag. (1854: 118)
= Marasmtus restnosus Peck (1883: 181) nom. nov. for
M. decurrens Pk.
Marasmius resinosus var. niveus Peck (1903: 38)
= Marasmius restnosus var. candtdtssimus Peck (1905: 40)
nom. nov. tlleg. for M. resinosus var. niveus Pk.

WU va

PILEUS: 4-15(-19) mm wide, convex becoming plano-convex to plane,
usually depressed centrally, occasionally umbilicate, rarely
subumbonate, white to yellow white, opaque to vaguely striate
marginally becoming obscurely corrugated-striate on some and sometimes
obscurely concentrically ridged, dry to tacky or slightly viscid,
micaceous when dry; edges incurved at first, uneven with age; context
tough-pliant, concolourous; odor and taste not distinctive.

LAMELLAE: adnate to subdecurrent or arcuate-decurrent, whitish,
moderately narrow, close, often becoming forked in places or
developing ladder-like anastomoses; edges crenulate, in some specimens
beaded with resin. STIPE: 12-50 nm long, 0.5-1 mm wide, equal or
slightly enlarged above, tough and pliant, drying to a cormeus
texture, whitish, varying from glutinous to tacky or dry and
glistening from resinous cells in scattered beads, fistulose, with a
silky radiating white basal disc or subiculun.

PILEAL EPICUTIS: a layer of polymorphic, suberect, later
decumbent and tangled cystidia ranging from cylindrical narrowly
clavate or capitate oleocystidia to narrow dendroid nonresinous forms
with some intergradation, 40-60 X 3-9.5um, hyaline, thin-walled, often
filled or partially filled with vacuolate to homogeneous oily contents
and covered with similar exudates which dry as slightly yellowish and
hardened masses. PILEAL TRAMA: obscurely duplex, with slightly
broader hyphae above, 5-15um diam., than below, 3-10um diam.; hyphae
subparallel, clamped, smooth, walls thin to slightly thickened,
faintly pseudoamyloid. LAMELLAR TRAMA: similar to pileal trama,
varying to slightly interwoven, more definitely pseudoamyloid.
CHEILOCYSTIDIA: abundant, forming a sterile edge, similar to the
cylindrical to narrowly clavate or fusoid oleopileocystidia,

55-65 X 7-7.5um. PLEUROCYSTIDIA: rare or absent, similar to the
cheilocystidia. BASIDIA: 19-22 X 5-5.2yum, narrowly cylindrical to
obscurely utriform, 4-spored, clamped. BASIDIOSPORES: 5.4-8.5 X
2.4-4.1(-4.5) um often being predominantly at one or the other end of
the range, mostly ellipsoidal or broadly cylindrical, inequilateral in
profile, prominently apiculate, smooth, hyaline, thin-walled, amyloid.

STIPE HYPHAE: parallel, 3-5ym diam. in the cortex, up to 10um diam.
in the medulla, mostly prominently pseudoamyloid in the medulla, walls
uneven and slightly thickened. CAULOCYSTIDIA: similar to the
pileocystidia, often clustered. BASAL MYCELIUM: interwoven, 2-3um
diam., smooth, inamyloid, clamped, hyaline, walls refractive and
slightly thickened.

HABITAT, HABIT and SUBSTRATES: scattered to subcespitose on leaf
litter, small twigs or bits of wood and woody fruits or husks from
Rhododendron, Quercus, Fagus, Castanea, Carya and sometimes Pinus when
mixed with Quereus in eastern hardwood forests.

COLLECTIONS EXAMINED: (only collectors initials cited after first
full citation): CANADA: Ontario: London, Sept. 27, 1896, Aug. 18,
1897, Sept. 8, 1915, J. Dearness (DAOM); and probably London area,
sept. 11, 1903 and October 23, 1926, J.D. (DACM); Toronto, July 12,
1932, H.S. Jackson (DAQM 50112, ex TRIC 3381). U.S.A.: Georgia:
Rabun Co., Rabun Bald, Sept. 3, 1947, Walters 153 (MICH). J2linots:
River Forest, bank of Desplaines R., July 1, 1902, E.T. & S.A. Harper
542 (F). Kentucky: Harlan, Sept. 4, 1916, C.H. Kauffman (MICH).
Michtgan: Lenawee Co., Cleveland L., Omsted State game area,

pepe. 9, 1970, W. Patrick (M. Gilliam 939; MICH); Livingston Co.,
George Reserve, July 27, 1970, J. Williams (M.G. 834; MICH), Pinckney,
Bide 195) ,eAstH. omith 68235 (MICH), Oakland '\Co-,, Proud L., Oct. 10,
PI7O 70M. G.) 992) (MICH), Haven’ Hill, Aug. 8, 1972, A.H.S. 81375 (MICH);
MiesrtLemaweCOl. me Ali ALDOL, sUULV 920, 0.912, C.HaKe, OCt. /,,1931, (A-HsSey
irae 935 8A... 11699, Aug. 922, 1937,,A.Heo. /1/5, Aug. 11, 1960,
Mensa enol5/3, July 11, 1970, C. Nimke (M.G. 693, 701), July 30, 1970,
C. Nimke 49 (MICH), Gorman L., Aug. 20, 1972, A.H.S. 81620 (MICH),
Halfmoon L., July 26, 1970, S.J. Mazzer 6186 (MICH), Manchester,

mtv 4,9 1935, A.H.S. 1470 (MICH), Sharon Hollow, Aug. 4, 1960, R.L.
Shaffer 2513, Aug. 19, 1970, M. Gilliam 929, Sept. 16, 1970, M.G. 952
(MICH), Silver L. area, Aug. 22, 1960, R.L.S. 2646 (MICH), Waterloo
Rec. area, Sept. 3, 1968, F. Hoseney 1094, July 10, 1970, F. Hoseney
Mec mol), uly 22,.1970, M.G../86, July 2,,1971, F.H. 1800 (MICH),
Winnewana, July 11, 1970, M. Gilliam 648 (MICH). Minnesota: Rice
Co., Wheeling Two., Nerstrand State Park, Aug. 29, 1965, M.G. Weaver
1243 (MICH). Worth Carolina: July 11, 1924, J.V. Couch 7380 (MICH),
Swain Co., Flat Cr., Aug. 1, 1937, L.R. Hesler & A.J. Sharp (MICH).
New York: Albany, rural cemetery, July-Aug. 1872, C.H. Peck [type of
M. decurrens] (NYS); East Worchester, July, C.H.P. (NYS); Forestburgh,
Sept. 1874, C.H.P. [type of A. rhododendri] (NYS); Genesee Co., Bergen
Swamp, Sept. 4, 1972, H.S. Vishniac (MICH); Greenbush, Aug., C.H.P.
(NYS); Ithaca, Aug. 11, 1904, C.H.K. (MICH), Aug. 1935, H.C. Beardslee
Jr. 35074 (MICH); Port Jefferson, Aug. 6, C.H.P. [type of M. restnosus
var. niveus] (NYS), Schuyler Co., Hector land rice area, Aug. 19, 1972,
feSeVea(MICH). Ohio: Lane; July 13, 1922, H.C.B. Jr. (MICH); Portage
Co., West Branch State Park, July 8, 1972, M. Gilliam 1493 (MICH) .
Pennsylvania: Mt. Gretna, Sept. 5, 1926, C.H.K. (MICH) ; Media,

June 29, 1940, P.M. Rea & Woodbury (MICH). Tennessee: Great Smoky
Mts. Park, Cades Cove, Aug. 18, 1938, A.H.S. 10374, Indian Gap,

Aug. 29, 1938, A.H.S. 10625, Aug. 7, 1942, L.R.H. 14476, Laurel falls
ert eA 5, 01936,0A.H.S. 9909 (MICH).

Restnomycena rhododendri was first described by Peck (1872)
as Marasmius decurrens. He apparently collected a similar fungus in

LSS

Figs. 1-5. Restnomycena rhododendrt (DAOM, ex Dearness Oct. 23,
1926). 1, basidiospores. 2, basidia. 3, pileocystidia.
4, caulocystidia. 5, cheilocystidia.

WMHs)

Greenbush in 1869, #20, but lost the specimen before describing it

(J. Haines, pers. comm.). Peck's validating description of

A. decurrens indicated a dark taxon with a grayish or tawny pileus and
a gray stipe. These colours are not apparent on the type of

A. decurrens, microscopically or macroscopically, possibly indicating
that more than one taxon was involved when the description was
written. However, as the name is now lectotypified by Gilliam (/.c.),
it applies to the pale form described in this paper. Peck (1883) not
Saccardo (1887) as is often cited, later proposed the new name

M. resitnosus for M. decurrens which was a later homonym of

M. decurrens Mont. However, Peck (1875) had in the meantime
unknowingly described it a second time as Agaricus rhododendrt which
is, therefore, the earliest valid and legitimate name. His variety
niveus is herein treated as being con-varietal with the type variety
as it is now typified. The proposal of the new name M. restnosus var.
candtdisstmus to replace M. restnosus var. niveus is superfluous under
present nomenclatural rules, but it should be kept in mind that Peck
was following the Rochester Code.

Smith (1947) was the first to recognize that Agartecus rhododendrt
and Marasmius restnosus were congeneric, although he reported the
spores of A. rhododendrt to be smaller, 3-4.5 X 2.5um, than for
M. restnosus. Bigelow (1970) treated the two as a conspecific but
offered no evidence. In our studies spores measuring 6.5-7.2 X
3-3.2um were found on the type of A. rhododendri which in other
respects is typical for M. restnosus. Thus the two are treated as
conspecific. Hesler's (1959) report of inamyloid spores for the type
of M. restnosus is evidently in error as we concur with Gilliam (1976)
that amyloid spores are present.

Restnomycena rhododendrt appears to be restricted to the eastern
deciduous forest where it occurs on litter of the dominating hardwood
trees. It is the largest of the Restnomycena species and the one most
commonly collected.

Restnomycena kalalochensts (Smith) Redhead & Singer, comb. nov.
Figs. 6-10.
= Mycena kalalochensts A.H. Smith (1947: 99)

"Pileus 3-8 mm. broad, convex, remaining broadly
convex, margin incurved at first, spreading in age, chalk
White and appearing pruinose under a lens at first,
glabrous and uneven in age but remaining chalky, slightly
sulcate at maturity, not hygrophanous; flesh membranous
and pliant (but not reviving), odor not distinctive, taste
not recorded; lamellae adnate, broad, distant, 10-12 reach
the stipe, two tiers of lamellulae, white over all, edges
pruinose; stipe 3-7 mm. -long less than 0.5 mm. thick,
equal or the base flanged slightly, strigose, the remainder
pruinose like the pileus, chalky white over all."

Smith (1947: 99-100).

PILEAL EPICUTIS: an erect to suberect tangled turf of
polymorphic cystidia, 25-50 X 3-8um, varying from cylindrical to
narrowly clavate, usually capitate to subcapitate, often secondarily
septate, scantily resinous oleocystidia to variously branched to

narrow dendroid and antler-like forms, which lack resinous exudates
or contents, walls thin, hyaline, smooth, clamped basally. PILEAL
TRAMA: somewhat duplex with slightly broader hyphae above, 5-15um
diam., than below, 5-10u diam.; hyphae faintly pseudoamyloid
initially but becoming more intense after several days in permanent
Hoyer's-Melzer's reagent, clamped, smooth, thin-walled, subparallel.
LAMELLAR TRAMA: hyphae similar to the pileus trama hyphae.
CHEILOCYSTIDIA: abundant, forming a sterile edge, less variable than
the pileocystidia, mostly narrowly cylindrical to slightly clavate,
occasionally forked or branched, occasionally with an apical
finger-like elongation, 31-48 X 5-6.5um, scantily resinous. BASIDIA:
24-25 X 7.5-8.0um, clavate to obscurely utriform, 4-spored, clamped.
BASIDIOSPORES: 7.8-10.8 X 3.8-4.8um, ellipsoid to obscurely fusoid
or occasionally obovoid, inequilaterally flattened in profile,
smooth, thin-walled, amyloid, hyaline, with a prominent apiculus.
STIPE HYPHAE: parallel, 5-15um diam., smooth, pseudoamyloid,
hyaline, with the broadest hyphae in the medulla. CAULOCYSTIDIA:
similar to the pileocystidia but only scantily resinous like the
cheilocystidia. BASAT, MYCELIUM: not studied.

HABITAT, HABIT and SUBSTRATES: Scattered on litter of Rubus, Alnus
rubra Bong. and grasses in the coastal forest zone of western North
America.

COLLECTIONS EXAMINED: CANADA: Brtttsh Columbia: North Vancouver,
Capilano Canyon, Oct. 8, 1973, S.A.R. #AS 9 (DAOM 166524), Vancouver,
Warvemts Cpe uuner2, LOAN S.Avk. 3425 (UBC) .0 UsoeAas) SCaLtrornia:
Humbolt Co., Big Lagoon Park, Dec. 18 & 19, 1956, A.H.S. 56787,
50033,5 50823, Prairie Cr. Park, Dec.99, 1956, A.H-S. 56520 (MICH).
Oregon: Seaside, Sept. 21, 1944, W. Gruber & A.H.S. 19020 (MICH).
Washtngton: Jefferson Co., Kalaloch, April 30, 1939, A.H.S. 13035
[type] (MICH).

A.H. Smith (1947) reported slightly larger spores, 8-11 X 5-6um,
an inamyloid lamellar trama and a lack of incrustations on the
pileocystidia. In our studies resinous exudates and dextrinoid
tramal tissues were found on the type although the iodine reaction
was not as prominent as that for the stipe. In addition, the
resinous exudates tend to become clarified and dissolve in Melzer's
reagent more so than in KOH 3% aqueous sol.

Resinomycena kalalochensts differs from R. rhododendrt by
distant lamellae, smaller basidiomes, larger spores, and more scant
resinous exudates, in addition to the differences in geographic range
and to a lesser extent substrates.

Restnomycena montana Redhead & Singer, sp. nov. Figs. 11-14,26.

Pileus 3.5-5.5 mm latus, convexus vel depressus, candidus.
Lamellae arcuatae, candidae. Sttpes 9-23 mm long., 0.4-0.6 mm
erassus, candidus. Sporae 8-9.3 X 2.8-4.5um, amylotdeae,
elltpsotdeae. Ptleocysttdia 45-67 X 2.5-7\m, restnosa, clavata vel
dendrotdea ut nonrestnosa. Chetlocystidta 42-52 X 6-6.2um, restnosa,
elavata vel strangulata. Tramate pseudoamylotdeo.

| |

; 10
Figs. 6-10. R. kalalochensts (DAOM 166524). 6, caulocystidia. c
7, basidiospores. 8, cheilocystidia. 9, basidia. 10, pileocystidia.

160

Figs. 11-14. FR. montana (DAOM 178214). 11, basidia.
12, pileocystidia. 13, cheilocystidia. 14, basidiospores.

Lot

Holotypus: DAOM 178215. Donald Station, Rocky Mt. trench, B.C.,
Canada, Sept. 21, 1980, S.A. Redhead 3977.

PILEUS: 3.5-5.5 mm wide, convex and slightly depressed
centrally, vaguely striate to chalky white, micaceous, obscurely
corrugated-striate, developing greyish casts when dry; context white,
pliant; odor not distinctive. LAMELLAE: arcuate decurrent,
moderately spaced, white, with micaceous edges; lamellulae in 2 tiers.
STIPE: 9-23 mm long, 0.4-0.6 mm wide, equal, cargilaginous or
tougher, white, finely powdered overall.

PILEAL EPICUTIS: a suberect to tangled turf of polymorphic
cystidia 45-67 X 2.5-7um, varying from a majority of elongated clavate
oleocystidia to filiform or dendroid nonresinous elements. PILEAL
TRAMA: obscurely duplex, with slightly broader hyphae above, 3-llum
diam., than below 3-5yum diam.; hyphae subparallel, faintly
pseudoamyloid at first, smooth, clamped, thin to slightly
thick-walled, hyaline. LAMELLAR TRAMA: hyphae similar to the lower
pileal trama hyphae. CHEILOCYSTIDIA: abundant, forming a sterile
edge, 42-52 X 6-6.2um, narrowly clavate to cylindrical and slightly
strangulate, occaSionally with a short apical elongation, with scant
to abundant resin exudates. BASIDIA: 20-21 X 6.8-7.2um, clavate
subcapitate, 4-spored, clammed. BASIDIOSPORES: 8-9.3 X 2.8-4.5um,
cylindrical to narrowly ellipsoidal, slightly inequilateral in
profile, amyloid, smooth, hyaline, with a prominent apiculus. STIPE
HYPHAE: parallel, pseudoamyloid especially on the slightly broader
medulla hyphae, 4-13um diam., smooth, thin to slightly thick-walled,
clamped. CAULOCYSTIDIA: scattered to abundant and often clustered,
similar to the pileocystidia. BASAL MYCELIUM: not studied.

HABITAT, HABIT and SUBSTRATES: scattered to gregarious on coniferous
needles, cone debris and small twig fragments in western Subalpine,
Coastal and Columbian coniferous forests of North America.

COLLECTIONS EXAMINED: CANADA: Brittsh Columbta: North Vancouver,
Mt. Seymour, 3500' alt., Sept. 17, 1973, S.A.R. #AJ 3 (DAOM 178218);
Sept. 22, 1973, S.A.R. #AI 24 (DAOM 178214); Columbia R. valley in
Rocky Mt. trench at Donald Stn. on Hwy. 1 crossing of river,

Sept. 21, 1980; S.A.R. 3977 [type] (DAOM 178215). U.S.A.:
Washington: Lower Elwha, July 3, 1939, A.H.S. 14732 (MICH).

Restnomycena montana has longer spores than Rk. rhododendrt,
occurs on coniferous debris and usually develops a greyish cast on
drying. Geographically the two are widely separated. Resinomycena
kalalochensis differs by its more capitate and shorter pileocystidia
Which are often secondarily septate and scarcely resinous. Also the
latter has smaller basidiomes which have not been observed to develop
a grey cast and it occurs on angiosperm debris.

Restnomycena acadtensts Redhead & Singer sp. nov. Figs. 20-25.

Pileus 2-10 mm latus, convexus denum depressus, candidus.
Lamellae arcuate, ecandidae. Stipes 10-40 (-70) mm long., 0.2-0.8 mm
crassus, candidus. Sporae 9.4-12.8 X 4-5um, anyloideae, elltpsotdeae.
Ptleocystidta 30-58 X 3-9um, restnosa, clavata vel nonrestnosa ut

162

dendrotdea. Chetlocystidta 38-40 X 6-8um, aeque ptleocysttdia.
Tramate pseudoamylotdeo.

Holotypus: DAOM 166073. Kouchibouguac Natl. Park, N.B., Canada
Sept. 24, 1977, J.E. & S.A. Redhead 2558.

PILEUS: 2-10 mm wide, convex becoming depressed to cyathiform,
translucent-striate but readily becoming opaque on partial drying,
white to whitish, occasionally tinted fawn on the disc, rugose with
age, moist to dry; edges incurved initially; context white, pliant;
odor and taste not distinctive. LAMELLAE: arcuate-decurrent,
becoming more distinctly decurrent, white, subdistant to moderately
spaced; edges crenulate and often beaded; lamellulae in 2 tiers.
STIPE: 10-40 (-70) mm long, 0.2-0.8 mm wide, equal or tapering up,
white or with ochreous to rosy buff tints basally, glutinous when wet,
powdered when dry, often attached to the substrate by a small
radiating silky white mycelial pad.

PILEAL EPICUTIS: a sSuberect turf and later repent tangle of
polymorphic cystidia 30-58 X 3-9um, varying from short to long clavate
oleocystidial forms to narrow filiform to dendroid nonoleocystidial
forms with some intergradation, thin-walled, smooth, hyaline, with
copious to moderate resinous exudates. PILEAL TRAMA: obscurely
duplex with the broadest forms in the upper half, hyphae subparallel,
5-llum, smooth, hyaline, faintly pseudoamyloid at first, clamped.
LAMELLAR TRAMA: hyphae mostly parallel, 3-5um diam., pseudoamyloid.
CHEILOCYSTIDIA: abundant forming a sterile edge, similar to the
pileocystidia but rarely of the branched forms, mostly clavate,

38-40 X 6-8um. BASIDIA: 20-32 X 7-8um, (2-)4-spored, clavate,
clamped. BASIDIOSPORES: 9.4-12.8 X 4-5yum, ellipsoidal to cylindrical
or obscurely reniform to pip-shaped, inequilaterally flattened in
profile, with a prominent apiculus hyaline, white in mass, amyloid.
STIPE HYPHAE: 3-10um diam., with narrower hyphae in the cortex than
in the medulla, strongly pseudoamyloid, smooth, parallel, hyaline,
clamped. CAULOCYSTIDIA: usually clustered, similar to the
pileocystidia but less often with long pedicels and more often
secondarily septate. BASAL MYCELIUM: slightly interwoven,
agglutinated, 3-10um diam., pseudoamyloid.

HABITAT, HABIT and SUBSTRATES: scattered on small broken pieces of
woody angiosperm debris in predominantly Acadian coniferous forests of
larch or spruce, fir and pine.

COLLECTIONS EXAMINED: CANADA: Wew Brunswick: Kouchibouguac Natl.
Park, Aug. 11, 1977, R.L. Milikin (DAOM 169619), Sept. 24, 1977,
J.E. & S.A.R. 2558 [type] (DAQM 166073), Oct. 9, 1978, S.A.R. 2753
(DAOM) .

Restnomycena acadtensts can be distinguished from R. rhododendri
by the larger spores, more distant lamellae and the more flaccid
flesh. Restnomycena kalalochensts differs by its small stature, its
more erect turf of capitate and scarcely resinous, secondarily
septate pileocystidia and its geographic range. Restnomycena montana
differs by its smaller usually narrower spores, smaller basidia, a
coniferous substrate and its geographic range.

The glutinous to viscid stipes are a result of copious resinous

163

exudates. These dry down to hardened masses in less humid situations
creating a powdered appearance to the stipes. In herbarium specimens
these masses become yellow with age.

Restnomycena brunnescens Redhead & Singer, sp. nov. Figs. 15-19.

Pitleus 6 mm latus, planoconvexus ut depressus, rugulosus,
brunneus. Lamellae adnatae vel subdecurrentes, bubalineae vel
ochraceae. Sttpe 20 mm tlong., 0.5 mm crassus, concolorus. Sporae
6.5-8 X 3.3-4um, amylotdeae, elltpsotdeae. Ptleocystidia 55-90 xX
3-1lum, restnosa, clavata vel strangulata, vel nonrestnosa ut
dendrotdea. Chetlocystidia 36-50 X 10-14um, resinosa, clavata vel
capttata, rarts nonrestnosa ut dendrotdea. Tramate pseudoamylotdeo.

Hototypus: DAOM 165884. Kouchibouguac Natl. Park, N.B., Canada,
July 13, 1977, J.E. & S.A. Redhead 2336.

PILEUS: 6 mm wide, plano-convex and depressed centrally, rugose,
tacky, scarcely translucent on edges, greyish sepia centrally, fulvous
with ochreous to buff margins; context membranous, thin, concolorous
above; odor not distinctive. LAMELIAEF: adnate to subdecurrent,
moderately spaced, buff to ochreous with whitish crenulate edges,
sometimes forked; one tier of lamellulae. STIPE: 20 X 0.5 mm,
concolorous with the pileus with the darkest pigments below, with a
striated powdered appearance from dried resin, micaceous, attached to
the substrate by a radiating white mycelial pad.

PILEAL EPICUTIS: a layer of tangled polymorphic cystidia
55-90 X 3-llum, varying from clavate to fusoid often strangulate
oleocystidia exuding copious resinous exudates to filiform often
branched dendroid nonresinous forms. PILEUS TRAMA: duplex, with
hyphae 10-15yum diam. subparallel, thin-walled, pseudoamyloid,
yellowish brown and prominently incrusted with brownish resin-like
materials in flat patches above, 5-10um diam., hyaline, walls thin or
pronounced below. LAMELLAR TRAMA: hyphae similar to those of the
lower pileus trama. CHEILOCYSTIDIA: abundant, forming a sterile edge
which on drying is a continuous hardened resinous line, 36-50 X
10-14um, clavate and prominently capitate, hyaline, thin-walled.
BASIDIA: 27-29 X 5-6um, narrowly clavate (2-)4-spored, clamped.
BASIDIOSPORES: 6.5-8 X 3.3-4yum, ellipsoid to obscurely obovoid or
fusoid, hyaline, smooth, thin-walled, amyloid, with a prominent
apiculus. STIPE HYPHAE: parallel, 3-5ym diam. in the cortex with
pigments similar to the pileus tramal hyphae, 10-15um diam. in the
medulla, pseudoamyloid, clamped. CAULOCYSTIDIA: clustered, similar
to the cheilocystidia but less swollen apically, occasionally forked,
intermixed with a few dendroid filiform elements. BASAL MYCELIUM:
not studied.

HABITAT, HABIT and SUBSTRATE: solitary on fallen leaf of Rhododendron
canadense (L.) Torr. in a dense Acadian coniferous forest.

COLLECTION EXAMINED: CANADA: WNew Brunswick: Kouchibouguac Natl.
Park, July 13, 1977, J.E. & S.A.R. 2336 [type] (DAOM 165884).

Resinomycena brunnescens is readily distinguished from all other
species in the genus by its darkly pigmented pileus and stipe. The

165

Figs. 15-19. R. brunnescens (DAOM 165884). 15, basidia.
16, cheilocystidia. 17, caulocystidia. 18, basidiospores.
19, pileocystidia.

167

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Figs. 20-25.

168

incrusted hyphae responsible for this darkened appearance are very
distinctive microscopically compared to the nonincrusted hyphae in
other Restnomycena species. In addition, R. brunnescens is
characterized by very prominently capitate cheilocystidia which are
inflated more than in any of the other species.

As noted above, Peck's original description of Marasmtus
decurrens is of a brown fungus. The existence of a brown species of
Restnomycena in eastern North America suggests that Peck may have
collected both R. brunnescens and R. rhododendrt. However, this
cannot be confirmed now.

GENERAL DISCUSSION

The three most probable generic depositories for Resinomycena if
the taxon is not recognized as a distinct genus are Hydropus, Mycena
and Baeospora, there being a foregone conclusion that the species are
excluded from the genus Marasmtus because of amyloid spores. The
genus Baeospora as it was defined in The Agartcales tn Modern Taxonomy
(Singer, 1975) represents a small cohesive genus characterized by a
number of characters, but in particular, small amyloid spores,
inamyloid tramal tissues and crowded lamellae. Restnomycena
rhododendrt is the species most similar to a Baeospora in that it has
close lamellae and small amyloid spores, however, the tramal tissues
of the pileus and stipe are pseudoamyloid unlike any Baeospora
species. Also the massed large oleocystidia mixed with dendroid
elements on the pileus form a structure unlike any epicuticular
formation found in Baeospora, including B. pallida. When all species
of Resitnomycena are taken into account, species such as R#. acadtensts
and R. kalalochensits, exhibit characters not found in Baeospora such
as distant, slightly decurrent lamellae and large spores. As a result
of these combinations of characters Restnomycena cannot readily be
incorporated into Baeospora.

The genus Hydropus as defined by Singer (1975) contains species
having numerous pileo-, cheilo- and caulocystidia and in some species
pseudocystidia with oily contents. The spores can be amyloid and the
tramal tissues can be weakly pseudoamyloid. Again the abundant
Oleocystidia on the pilei, stipes and lamellae of Restnomycena
species, not found in Aydropus, in combination with intermixed
dendroid elements form a tissue not found in Aydropus and clearly
separate the genera.

The genus Mycena is treated as delimited earlier (Singer 1975).
It includes species with amyloid spores, pseudoamyloid tramal tissues
and oleocystidia as does Hydropus. The majority of Mycena species
have an epicutis consisting of a repent layer of diverticulate narrow
hyphae. Deviations occur in some sections of the genus where smooth
repent filamentous hyphae replace the diverticulate elements. Those
species which had smooth epicuticular hyphae bearing obtuse cystidia
were removed to the genus Hydropus. The most aberrant type of
epicutis for a species admitted to Mycena by Singer (1975) is found in
Mycena rorida (Fr.) Kuhner and its allies. ‘These species have a
hymeniform layer of sphaeropedunculate cells and, apparently missed by
previous authors, in M. roritda, intermixed oleocystidia. Previously

(Singer 1975), it was indicated that these species were related to
Hydropus and might need to be removed from Mycena. However, the
strongly pseudoamyloid tramal tissues and a gelatinized stipe cortex
weighed in favor of retaining the species in Mycena. Here again, the
well developed layer of usually clavate oleocystidia and narrow
dendroid elements found in Restnomycena is not typical of any section
of Mycena and represents a large divergence from a typical Mycena type
of epicutis. Furthermore, the typical Mycenae have characteristically
broad (broader than Resinomycena) hypodermal and tramal hyphae; the
hyphal cells often being short and voluminous. In a less restrictive
delimitation of Mycena such as by Smith (1947) or Kuhner (1938),
Restnomycena would be included.

From the above discussion it can be seen that Baeospora,
Hydropus and Mycena are open to emendation to encompass Resinomycena
species but only by introducing a species group having an unusual
combination of characters peripheral to the core of the genera
involved. Instead the proposal to recognize a distinct genus seems
more justified. Restnomycena species are obviously closely related to
each other. They are readily distinguished from Baeospora, Hydropus
and Mycena by the abundance of resinous oleocystidia intermixed with
dendrohyphidia. The segregation of Restnomycena species maintains the
integrity of the older genera, leaving them less heterogeneous and,
therefore, more recognizable. There is no doubt that Resinomycena is
closely related to Baeospora, Mycena and Hydropus. Obviously one
solution would be to merge all three genera and add Resinomycena.
Again there seems to be more merit in recognizing each of these genera
including Restnomycena.

Finally, Marasmius rhododendrt Singer (1947, 1936), a species
described from Spain in the subalpine zone of the Pyrenees on
Rhododendron ferrugineum L. is probably congeneric. ‘The type (F) now
consists of stipes only. Spores obtained from the type were amyloid
although weakly in some and up to 12.5um long. The stipe hyphae are
pseudoamyloid and in an outer layer more or less gelatinized (the
stipe was described from fresh material as somewhat shiny and
viscidulous). They bear cystidia with dried incrustations similar to
that found on Restnomycena species. In the original description
(1936) yellowish incrustations were mentioned on the lamellar edges of
vesciculose to ampulliform cheilocystidia ("echinulatis" possibly
being an error in the 1947 description unless referring to dendroid
elements).

The exact nature of these cystidia and the pileipellis could not
be restudied in the damaged type collection and the identity of the
species remains in doubt. The weakly developed collar is not found
on the presently recognized Restnomycena species. This species
should be restudied from fresh materials.

ACKNOWLEDGEMENTS

We thank Drs. J. Haines (NYS), R.J. Bandoni (UBC), R.L. Shaffer,
R. Fogel and A.H. Smith (MICH) for the use of herbarium materials and
facilities. Drs. J. Ginns and J. Ammirati offered appreciated reviews
and Mr. K.W. Spicer provided technical assistance. Parks Canada
supplied collecting permits or facilities in New Brunswick and British
Columbia and the Agriculture Canada research station, Kamloops,

170

provided transportation in 1980. Mrs. M. Meredith typed the final
manuscript.

LITERATURE CITED

Bigelow, H.E. 1970. Omphaltna in North America. Mycologia
G2 spelen 3 2

Gilliam, M.S. 1976. ‘The genus Marasmtus in the Northeastern United
States and adjacent Canada. Mycotaxon 4: 1-144.

Hesler, L.R. 1959. Southeastern Agaricales, IV. J. Tenn. Acad. Sci.
34: 167-171.

Kuhner, R. 1938. Le genre Mycena. Encyl. Mycol. 10: 1-710.

Montagne, J.F.C. 1854. Cryptogamia Guyanensis seu plantarum
cellularium in Guyana gallica annis 1835-49 acl. Leprieur
collectarum enumeratio universalis. Ann. Sci. nat. bot.,
sér. 4, 1: 91-144.

Murrill, W.A. 1916. Omphalopsts. N. Amer. Flora 9(5): 310-318.

Peck, C.H. 1872. Report of the botanist. Ann. Rep. N.Y. State Mus.
Nat. Hist. 24: 41-108.
. 1875. Report of the botanist. Ann. Rep. N.Y. State Mus.
NatwHist. ruc] s/o iG.
~ L883.) -Orders]16.).Fungive Dp. 1/4235 eines eDay ame
plants of Buffalo and its vicinity. Bull. Buffalo Soc. Nat.
SCT esa (4): 65-279.
. 1903. Report of the state botanist. N.Y. State Mus.
Bullvo1ce Lolo.
. 1905. Report of the state botanist. N.Y. State Mus.
Bull. ~94:°19=50:

Saccardo, P.A. 1887. Sylloge Fungorum. V. 1-1144. Patavia.

Singer, R. 1936. Notes sur quelques Basidiomycétes. Rev. Mycol.
(N.S) RC 2) seb =84
. 1947. Champignons de la Catalogne. Espéces observées en
1934. Collectanea Bot. 1(3): 199-246.
. 1975. The Agaricales in Modern Taxonomy. 3rd. ed.
J. Cramer, Vaduz.

1977. Interesting and new species of basidiomycetes from

Ecuador II. Nova Hedw. 29: 1-98.

Smith, A.H. 1947. North American species of Mycena. Univ. Mich.
SCI eSer. we liseli

MYCOTAXON

VO Lee el ee NOSSEL PSG Dp Dic guile 17 4 Api JUneeLo6

SINOTERMITOMYCES, A NEW GENUS OF AMANITACEAE
FROM YUNNAN, CHINA

ZANG MU

Kunming Institute of Botany, Academia Sinica, Kunming, Yunnan, China

SUMMARY

Detailed analysis has led the author to propose the termito-
philous Sinotermitomyces as a new genus, including two new
species, S. cavus Zang and S. carnosus Zang. Both were col-
lected in Yunnan, China. Type specimens of the two new spe-
cies are deposited in the Herbarium of the Kunming Institute
of Botany, Academia Sinica (HKAS).

The new genus Sinotermitomyces consists of two new species growing
on termite combs. These edible fungi are distributed in subtropical
and tropical regions of southwestern Yunnan. In Yunnan, profession-
al collectors gather both Termitomyces and Sinotermitomyces in the
rainy season from May to September, and dry preservation with salt
and vegetable oils are used for building up a year-round supply
(Cheo, 1942, 1948; Batra, 1979).

The genus Termitomyces in China is commonly found southward of
the Yangtze. The distribution of Sinotermitomyces is in forests of
the southwestern part of Yunnan. Specimens were collected by the
author under Pinus khasya Royle ex Cord., Pinus yunnanensis Fr.,
ande@astanopsisshystrixs (Hk. f.°6 Thoms.) 1A. DCs

SINOTERMITOMYCES Zang, gen. nov.

Pileus conicus vel campanulatus, subconvexus vel papillatus, siccus,
glaber vel scabridus. Lamellae subliberae, liberae vel adnatae, albidae.
Stipes centralis, cavus, aequalis vel sub-fusoideus, carnosus, coriaceus
vel fibrillosus. Annulus superus distinctus. Basidiosporae hyalinae,;
ellipticae vel obovatae, inamyloidea. Pleurocystidia ventricosa vel cy-
lindrica, verrucosa vel tuberculosa. Cheilocystidia oblonga vel cylindri-
Ca, laeviuscula vel verrucosa.‘ Fibulis nullis. Termiticola.
Typus generis: Sinotermitomyces cavus Zang.

Pileus conico-campanulate, subconvex and often with a prominent
papillate apex, dry, never viscid, glabrous or scabrous. Gills free
to almost free or adnate. Stipe central, often very long, variable
in. thickness, hollow, equal or. subfusiform, fleshy or leathery or
fibrillose. Annulus superus present, veil a pellicular veil that co-
vers the gills in young specimens. Basidiospores from globose to
ellipsoid, smooth, hyaline, inamyloid. Pleurocystidia present, cylin-
dric or some fusoid-ventricose, covered with wart-—-like outgrowths.
Cheilocystidia oblong to cylindric, smooth to warted. Clamp connec-
tions absent. The primordia develop in the holes of Nypogeous termi-

taria.

This new genus can be distinguished from the genus Termitomyces
Heim by the following characters: the small, subconvex pileus with
prominent papillate apex; presence of veil remains on stipe and
pileus; stipe leathery or fibrillose and hollow; pleurocystidia with
sturdy, short processes (Heim, 1942; Singer, 1949).

1. Sinotermitomyces CAVUS Zang, sp. nov. (FIGS. 1-7)

Pileus 1-2.5 cm., conicus vel subconvexus, siccus, glaber, primo flavi-
dus vel eburneus, demum fulvo-umbrinus vel rufo-—brunneus. Lamellae sub-
liberae vel adnatae, albidae vel eburneae, Stipes 20-30 cm. longus, 5-
12 mm. crassus, aequalis, cavus, tubulosus, coriaceus vel fibrillosus,
apice cicatricatus, sursam scabroso-furfuraceous, basim versus discoide-
us. Annulus superus distinctus.» Basidiosporae 2.4-5 x 3.5-9 pm, hyali-
nae, globosae, ovoideae, ellipsoideae, laeves, inamyloideae. Basidia
clavata, 15-25 x 10-14 pm, 4-sporigera. Pleurocystidia 12-20 x 25-45
pm, cylindrica vel clavata, verrucosa vel tuberculosa. Cheilocystidia
10-15 x 20-30 pm, oblonga vel cylindrica, tuberculosa. Fibulis nullis.

Hab. In sylvis praecipue Pinetis yunnanensi Fr. et Pinetis khasyae
Royle ex Cord. nido Termitidarum (Odontotermes) erumpente.

Yunnan: Tengchung County, Tuan-Tian village, La-ba-qing. On nest
of termites, in swampy grassland, under the pine woods above 2100 m.
alt., 8. Vill. 19802° Li’ Xing-jiange 11. (Typus,; HKAS +6533);) tengehung
County, Pu-Chuani-village,e17. Vil. 197908 Maw Xi—xiongiele com HKAcm40.i2.
4613); Shweli County, Deng Ga, in pine woods, 13. VIII. 1980. Li Xing-
jiang 22. (HKAS 6568); Mangshi. 10. VIII. 1980. Zang Mu 06545 (HKAS
6545).

Pileus 1-2.5 cm broad, conical or subconvex, dry, glabrous, surface
ivory white or pale yellow becoming yellowish-umber or reddish-
brown towards the center, glabrous. Gills almost free or adnate,
white to ivory white. Stipe 20-30 x 0.5-1.2 cm, equal, hollow, tube-
like, leathery to fibrillose, upwards often covered with bran-like
scales or scabrous and with a discoid base. Veil persistent as a
thick, membranous leathery annulus, which is striate above. Basidio-
spores 2.4-5 x 3.5-9 pm, hyaline, globose, ovoid to ellipsoid,
smooth, inamyloid. Basidia 15-25 x 10-14 wm, clavate, 4-spored.
Pleurocystidia 12-20 x 25-45 pm, cylindric to clavate, covered with
irregular wart-like outgrowths. Cheilocystidia 10-15 x 20-30 pum,
oblong to cylindric, roughened. Clamp connections absent.

2. Sinotermitomyces CARNOSUS Zang, sp. nov. (FIGS. 8-10)

Pileus 4-6.5 cm., conicus vel campanulatus, demum plani-convexus, sic-
cus, scabridus vel subtomentosus, primo albidus vel pallidus, demum cer-
vinus vel brunneus. Lamellae subliberae vel adnatae, albidae. Stipes
15-20 cm. longus, 1-2.5 cm. crassus, subfusoideus, carnosus, cavus, con-
colorous, pseudorhizophorus. Annulus membranaceus, superus instructus.
Basidiosporae 3-6 x 4.9-8 ym, hyalinae, globosae, ovoideae, ellipsoid-
eae, laeves, inamyloideae. Basidia 9.8-12 x 14-18 wpm, clavata, 4-spori-
gera. Pleurocystidia 10-18 x 24-48 pm, lageniformia vel fusiformia, exas-
perata vel verrucosa. Cheilocystidia 12-18 x 18-34 pm, rotunda, ovata,
oblonga, laeviscula vel exasperata. Fibulatae adsunt.

Hab. In sylvis praecipue Castanopsitis hystricis (Hk. f. & Thoms.)
A. DC. alt 1600-2100 m. nido Termitidarum erumpente.

Yunnan: Tsangyuan Autonomous County of Wa Nationality. On termites'
nest. 30. VIII. 1980. Zang Mu 06752 (Typus, HKAS 6752).

\
| :
tl

FIGS. 1-10, Sinotermitomyces spp.

(Jn. Scavus Zanes (HKAS 10533).50 91,
te Carpophores+ 2, .cheilocystidia;.3, pleurocystidia; 5, basidia and
basidiospores; 6, the discoid base of stipe. 8-10, 5S.
(HKAS 6752). 8, carpophores; 9,, cheilocystidia,
spores; 10, pleurocystidia.

carnosus Zang
basidia, and basidio-

173

174

Pileus 4-6.5 cm broad, conical, campanulate to plano-convex, dry,
scabrous or subtomentose, surface white or pale white to waxy yel-
low or brown. Gills almost free or adnate, white. Stipes 15-20 x
1-2.5 cm, subfusiform, hollow, more fleshy, uniform in colour, pseu-
dorhiza present. Annulus superior, fragments adhering to stipe. Basi-
diospores 3-6 x 4.9-8 ym, hyaline, globose, ovoid to ellipsoid,
smooth, inamyloid. Basidia 9.8-12 x 14-18 wm, clavate, 4-spored.
Pleurocystidia 10-18 x 24-48 ym, flask-shaped or fusiform, rough-
ened with wart-like outgrowths. Cheilocystidia 12-18 x 18-34 ym,
almost round, broadly oblong, almost smooth to roughened. Clamp
connections absent.

LITERATURE CITED

BATRA, L. R., (ed.). 1979. Insect-Fungus Symbiosis, pp. 142-145, 148-
153% Allanheld; Osmun & Cos, Montclair.

CHEOQ VC. eG. 1942. A study of Collybia albuminosa (Berk.) Petch, the
termite-growing fungus in its connection with Aegerita duthiei
Berk. Science Record 1: 243-248.

° 1948. Notes on fungus-growing termites in Yunnan, China.
Lloydia 11: 139-147.

HEIM, R. 1942. Les champignons des termitiéres. Nouveaux aspects d'un
probléme de biologie et de systématique générales. Rev. Sci. 80:
69-86.

SINGER, R. 1949. The Agaricales in modern taxonomy. Lilloa 22: 396-
398.

MYCOTAXON

ViOuLeweE tel Le aeN Of SD Dimer iio = OO April-June 1981

TYPE eo LUDT ES oLNe THE: POLYPORACHAR 13.
SPECIES SDESCRIBED IBY Js ho LEVELELE.

by
LEIF RYVARDEN

Botanical Laboratory
University of Oslo
P.O. BOX L045. blindern
Oslo 3, Norway

>. Us Meehan

Of the 145 polypores described by J. H. Léveillé

12 are accepted, 83 are synonyms, 6 names are in-
valid, 95 are an uncertain status, smost of them Ga-—
moderma sp.-, while 39 types were not. found. The
names of many of the missing types are accepted

as synonyms based on notes by previous mycologists.
abercombination Tyromyces dissectus is proposed.
epemracvver wpec eseand sl riachaptumeperrottetiiocare
described in detail.

The French mycologist J. H. Léveillé was very industri-
ous and described many fungi of which 145 are polypores.
His basis was the collections in the Paris herbarium be-
Sides which he made a journey to Leiden and was allowed to
take samples from the rich East-—Asian collections of Jung-
Dunn vanGrZovVlancer. Most, Ole hasecoLlectionsrvare ptoday jin
the Paris Herbarium (PC), but many of the polypore types
are also in the Leiden herbarium (L.). Isotypes are at Kew
(K), Stockholm (S), the Farlow herbarium (FH) and the Na-
tional Frunecus Collections (BRD).

Léveillé made very scanty labels, but often he fastened
his specimens to the sheet with a red silky band attached
both to the specimen and the sheet with a red lacquer. This
made it possible to study both sides of the specimen with-
out, loosening it’ from the sheet.

this red lacquershas in many cases made 1t possible? to
trace some of his types when label and specimen have been
separated, which was the case in many collections. Never-
theless, many of his types have been lost. However, both
Lloyd. (1912) and Bresadola (1916 & 1920) studied many of
his specimens before they disappeared and it is possible to
settle their identity. Léveillé described polypores in a
number of genera, and they are treated in the genus in
which Léveillé originally placed them. Within each genus

the species are placed alphabetically according to their
specific epithets. After the name there is a reference to
where it was published and an indication of the herbarium
in which the lectotype and eventual isotypes can be found.

The label is then cited in inverted commas and, if the
text indicates no type locality, this 18s added in brackets.
If theitype was found ‘to be 4 synonym this, is (indicated by
= followed by a correct citation of the name in question.

When the species has been accepted, it is cited in its
proper genus with a reference to a recent description; or
described in detail if no modern description seems to
exist.

DAEDALEA Fr.

D. aulaxina 1844:197. The type has not been found (Java).

= Lenzites vespacea (Pers.) Ryv. teste Bresadola
(1916:230).

Dem havica lous 31 OG IeCP Gr
"Java". = Lenzites acuta Berk.
D. fuliginosa 1844:199.
The type has not been found (Mauritius).
Den lurida, L644 OSS (PO® Vsotype dans).
Wav aie=s lerign tesa cuta sober.
Deamecrozonall oto dom ule
"Zollinger no 2060" (Java) = Lenzites acuta Berk.
D. plumbea 1846:142.
The type has not been found (Noveboracum = New York,

WShae
DY pruinosa 18845198 (PC) isotype in s)-
MING pacinico «= Lenzitesmacuta perk.

D. splendens 1844:197.

The type has not been found (Sumatra).
D. Violacea 18462142.

The type is apparently lost (Cuba).

FAVOLUS Fr.

Pee lOGa POsUsmlO ir OU ime CrCl.
"Manille" (The Phillipines). = P. phillipinensis Berk.
2p ah RAVES eee Aol (COG jee
"Bresil, Rio de Janeiro, Mars 1836" = F. brasilensis
CRs) br
F. granulosus 1863:286 (PC, isotype in K).
"Chachi alt. 2600 m, Sept...1860 Lindig 2919" -(Colombia).
= Panel lus pusillus - (bev. ) Duras.) & =Mlliler.
F. guadeloupensis 1846:144 (PC).
"Guadeloupe, M. 1‘'Herminier".
The type does not represent a true polypore and has
probably been fleshy and gelatinous when fresh. Today
it is resinous and horny. Pileus glabrous and pustulate,
dark brown, pores shallow and angular, 1-2 mm wide, dir-
ty to dark brown. Context dark brown and very dense. Hy-
phal system monomitic with clamps, variable in diameter,
in parts inflated, 2-8 um wide, but difficult to sepa-

rate. Spores and basidia not seen. The species is un-
known to me.

me junghuhnii 1844:202 (L, isotypes in PC & 8S).
DAGMUrUNCOStinmeinsulagbanvan’ ‘(rhe Phillipines) «= P.

hillipinensis Berk.
Meemultiplexelouu.203 (PCs) isotype in Kix S)-.

"Java". = F. spatulatus (Jungh.) Lév.
Pemepeltatus 18447203) (L, isotype in PC).

WZ0lvinger=no. 707" (Java). =F. brastlensis (Fr.) Fr.
Heetcener) Vo44; 202 (Lb).

"Sumatra". = F. spatulatus (Jungh.) Lév.
F. tenuissimus 1 wc Ue.s

Type not found (Mauritius).

GLOEOPORUS Mont.

G. leptopilus 1844:194.
Type has not been found (Surinam).

Gey pusililus 1844 :195.

I have not found the type, but the species has generally
been accepted as a poroid agaric. Modern descriptions can
be found in Singer (1945:224) who placed the species in
Dictyopanus, and in Burdsall and Miller (1978:85) who
transferred the species to Panellus.

HEXAGONIA Fr.

Hee plume ie 1 OA 1 99N (PC).
"Java, Blume". = H. tenuis (Hook.) Fr.

H. cingulata 1844:200.
The type has not been located (Java).

ie cyclophora 1846:143° (PC).
Mane te (Tahiti ).0.= sHeetenuits (CHook.) fr.
H. dregeana 1846:143 (PC).
NC. Bea. (Cape. of Good) Hope) Dreg. no. l/b.""= HH. tenuis
GHook eer.
H. glabra 1846:143 (PC).
"Bombay (India), Polydore Roux". = Lenzites acuta Berk.
H. molkenboeri 1844:260 (L).
"Java, Junghuhn". = Lenzites vespacea (Pers.) Ryv. The
synonymy was also noted by Bresadola (1916:231).
H. pulchella 1844:200 (L).
VJavalvmw= oH se tenlusm (HOOK... Er.
Pe itabacina, 1654-178 (PC, isotype in kK).
"Coll. Zollinger" (Java). = Cyclomyces setiporus (Berk.)
Pate

LENZITES Fr.

L. berkleyi 1846:122 (PC, isotypes in S & BPI).
"Grand bassin Saint Jean (Louisiana, USA) Mougeot no 36".
=i betulina (Pr.). Fr.

The type came from New York, but I have not found it.
Bresadola (1920-66) had apparently examined it and indi-
cated the synonymy given above. The cited specimen is
selected as neotype until the lectotype reappears.

LEG

Deectiiatae cud no I
No type has been found. No type locality was given.

L. guilleminiana 1846:122.
"Bresil meridional, Guillemin no. 1339" (PC). = Stipto-

BB llum erubescens (Berk.) Ryv.
junghubnii 1844:180 (PC).
"ad truncos Java" Léveillé's hand. "Pour moi identique
a halts betulina L. juvenile" Bresadola's handwriting.
=“eebetulina /Crrw Jr.
L. murina TOA 122) (PC te sO type. inno:
- ‘Java, Korthals". Selected as neotype. The type locality
is given assouma tra, DULL =nNO.such) CollecVion hasaueen
found. = Trametes menziesii? :(Berk.) Ryv.
Leamyriophyl lal 166322029 (PC):
Myovaes déeiJ. Triana 1651-57. -Novo (Granata’, (Colombia.
elegans (Fr.) Pat.
Ce Se LOUUT U7 OCPC Ya SOcLY pPeSmlibeNy sombre
"Java, ad truncos". = Lenzites vespacea (Pers.) Ryv.
L. platypoda Loud eeCO es CEO.
'"Manille (The Phillipines) ™.: = L. elegans (Fr. ) Par.
LOM Ceml USO or Loot scr Cnr
"Guadeloupe". = L. elegans (Fr.) Pat.

POLYPORUS | Er’:

P. apalus 1848:124.
No type has been found. (Rentilly, France).

Pemaonormasi bout 06".
The type has not been found (Java). = P. sanguinaria Kl.
teste Lloyd. Clo lO).

Plactleatus=! G462137 00? C eisotypesin BPr).
NZOdveinger no. 2055! “lava i=. trametes modes tamer re)
Ryv.

P. albomarginatus ©LG4W:.191 (PCy isotypes IniBPl kel).
No text on label, except the name written by Léveillé.
(Java). This is an accepted species in Pyrofomes as P.
albomarginatus (Lév.) Ryv. For a description, see Ry-
varden & Johansen (1980:529).

Pe Janisopilus loud oT EGRO.) Tsotypeswin suas worl).
"Java, Blume". = Coriolopsis sanguinaria (K1l.)’Teng-.

Pega pposttus el oloo iy Tip Geico t yneom Tien BE eo)
"Zoll. no. 19. Auf faulen Baumen in Wald bei Tjiboda
(Java) Mai 1842". This is an accepted species in Phelli-
nus' as P. appositus (Lév.) Pat. For a description, see
Ryvarden & Johansen (1980:141).

Poeatypus; Lond Cer c)

"Guadeloupe, M. L'Herminier". = Trametes modesta (Fr.) Ryv.
P. auriculaeformis 1844:194 (L, isotypes in BPI and “s)*

"Java". = Phellinus senex (Nees & Mont.) Imaz.
Peeplumelelold Looe Ty Wi soOtypes in bOrand Beh).

"Java, Zoll. no. 11" = Trametes menziesii (Berk.) Ryv.
P. bonplandensis 18463301, (PC. dsotynée an sBPL).

"Am equatoriale". = Trametes membranaceus (Fr.) Kreisel.

Pp. botryoides 16464120. (Sa sotypemin Beis)
"Wragg. orig. ex." (in Bresadola's hand) (North America).

= Globifomes graveolens (Schw.) Murr.

Paepracypusslcolo: 1? 7a(PChelsotypes ineBPT )’.
"Guadeloupe, M. L'Herminier, Février 1843". = Trametes
marianna (Pers.) Ryv.

fan coil imorphuse 16467133 7( PC).
"Voyage de M. Perville N.O. de Madagascar 1841."
Tp tee seangacceptedaspecves@inyPhellinus: aseP. vcalli=
morphus (Lév.) Ryv. For a description, see Ryvarden &
Johansen (1980:145).

Percaltochnrovice O44 Ol,
The type has not been found, and no type locality was
given.

Precandicanse| 005205. (PC. sOUY per ine hy.
"Tequendama (Colombia), alt. 2600 m, Sept 1856, Lindig
Noemecoe Te = ri chantumep i Pormis, (Free ins Kl.)) RYV.

Per candadulus 1846: 301" (PCRGisotype an’ Kk).
"Caesarodunum" (?) = Trametes cervina (Schw.) Bres. as
already noted by D. Reid on the sheet.

PB. chryseus 1846:301 (PC).
"Nouvelle Grenade (Colombia), M. Justin Goudot 1844.
ThVsm Svaneacceprednspecicceiteriel linus, as 72 .1cory—
seus (Lév.) Ryv. For a description, see Ryvarden & Jo-
hansenw 0603151").

P. cinerascens 1844:184.
The type has not been located (Java).

P. cineraceus 1846:139.
The type has not been seen by me, but Bresadola (1916:
223, as "cinereus") reports the species to be a synonym
Ol irametess hirsuta er.) bis

P. cohaerens 1846:132 (L, isotype in PC, BPI, FH and S).
"Zollinger no. 13" (Java). = Coriolopsis asper (Jungh.)
Teng. as already indicated on the sheet in herb. PC by
Bresadola.

Ree conrmertuswmlo4s Olu (CPG. Lsotype: Inv BPL).
"In truncos Java". = Coriolopsis asper (Jungh.) Teng.

P. eonnexus, 1846:135 (PC).
"Brasil, Rio de Janeiro. M. Gaudichaud 1831-33".
As already indicated by Bresadola (1920:68) the type
collection is mixed. One specimen is Coriolopsis cape-
rata (Berk.) Murr. while another is Phellinus gilvus
(Schw.)@ Pat. The description is so vaguel)-thatwit may
cover both specimens, and thus, the name has to be re-
jected.

Pe coniacenus, 16462137.
The type has not been found (Nelli Gherry, India). The
name is invalid as a homonym of P. coriaceus Endl. 1830.

Fe Cornmugacvus) 1o46ee41 36. (li)
"Java". Invalid name, being a homonym of P. corrugatus
Pers. 1826. The type of the invalid name is a specimen
of Coriolopsis asper (Jungh.) Teng.

Pee CONVO tUS! O44 MOOMCPC. sisotypes: in L,eSte NYoand BRL).
"Java, Junghuhn". = Trametes menziesii (Berk.) Ryv.

P.. cyathiformis 1844:181.
The type has not been found (Haiti).

Peademidofiiie boue:;02) (Ss, Pisotypesin BPl ands Ky).
"Frag. specimen orig. leg. Demidoff, comm. Patouillard".
This is an accepted species in Pyrofomes as P. demidof-
ble lev ve hOLlL ik sPoug. efor avdescription,) see Ryvar—
den (1978:399).

180

P. dermatodes, see Trametes dermatodes.
Paeaiii vatuse lout OlmLsy easotypeuines).
"java, ex religui Junghuhnii". = TraméetessmenZiesas
(Berk. ) eRY¥V.
P. disciformis, DOuts 1937
Thep type has not been found! (Mauritius).
Pevdissectusml S46. 13 9P¢PC)::
"Herbier du Chile, donné par M. Gay".

This isja Tyromyces species, and) the followings compi—
nation is proposed: Tyromyces dissectus (Lév.) Ryv.
comb. nov. Basionym: Polyporus dissectus Lév. Ann. Sci.
Nac emcees! VOU De So ml otbe

Fruitbody pileate, dimidiate with a contracted base,
ADOULE 2JDRxXE oabi cma Dasemabouc Wi cmuwidem ¢—Osnmatoick,
fragile. ‘Pileus applanate, margin fimbriate tovdastinct—
ly Split orelobeds surface white to dirtyelicn@ Drown,
smooth, azonate, finely velutinate to almost glabrous
in parts and weakly wrinkled radially. Pore surface
light brown, pores 4-5 per mm, entire to slightly in-
cised, tubes 2 mm deep, white. Context ochraceous, 1 mm
TOM Cutts |

Hyphal system monomitic, hyphae 2-5 um wide and with
clamps, partly gelatinized in 2.5 2% KOHes in) Melzerewith
walls about 0.5 um thick, sparingly branched. Some hy-
phae have a dense yellowish protoplasm, distinct both in
KOH and Melzer, 2-6 um wide and with scattered clamps,
these hyphae may be interpreted as oleiferous or gloeo-
pleurous. Cystidia and basidia not seen. Spores subcy-
lindrical to oblong ellipsoid, hvaline; thin-walled and
non—amyloid,. 4=-5.x 2-2.5 um. As long.as onlyethe: typeris
known, thesmacroscopical characteristics Nave fo, besused
with care.

This taxon may be the same as T. floriformis (Quel. )
Bond. & sing lheiremicroscoplecal eharacveristicsaare
1dentical and? the, fruitbodles arewsimilar. 1) flordior=
mis as collectedisins Buropel ie usualy morespurer wove.
buty themdarkerUcolours of tnemty pe Ole l.edLesecLusias
certainly due to bad preservation after it was collected
in Chile and then sent to France.

Pe dozyenus 1646 si23.
The type has not been found (Java).

Pe elatus 4] O46;129.
The type has not been found (Guadeloupe).

PV nex tensiswlo40e lo Um PCO):
"Guadeloupe, L'Herminier". This is an accepted species
in Phellinus as P. extensus (Lév.) Pat. For a descrip-
tion, see Ryvarden & Johansen (1980:158).

Pe. fastuosus, LO44eA 90 Nina teotype anbPl).
Jotnpapore!. Tne species Tseacceptedeunarhne: Linusewacgmr «
fastuosus (Lév.) Ryv. For a description, see Ryvarden &
Johansen (1980:159).

Peiruscelius, Lo54- 7 Flow Jeouypecwlneronanda la.
"Insul. Flores prope Balie". = Coriolopsis asper (Jungh. )
Teng.

P.tarusenus. O46: Lavoe POE soLyne in "ikl.
"Zollinger no. 1454" (Java). The type is a specimen of
what Berkeley later described as P. setiporus,:a species

181

which usually is placed in Cyclomyces. The type species
Cie bem obict isto. tUscusenr © 1033 making it illeriti-
mate to transfer Léveillé's name to Cyclomyces. However,
should anyone want to place the taxon behind Léveillé's
and Berkeley's names in a genus other than Cyclomyces,
then Léveillé's name has priority.
eeeeaudichaudii 1846p. 178) (PC, Wsotypes' in S & BPI).
"Singapore 1839". = Trametes menziesii (Berk.) Ryv.
ecavyanus 1 O46r1 ofr (PC)
Meniules Mem GlomGay..o30%.8 =—8Trametes marianna (Pers. )

Ryv.
P. gibberulosus 1846:139 (PC, isotypes in K & FH).
"Guyana, Batara Reidweg, Wegel no. 568". = Trametes vil-

HOSa HOOK) RY Ve

Pemecossypinus PG4sGe 124 (PC, isotype in’ BPI).
"Ad truncos in Voges (France)". = Tyromyces caesius
Chr Munim

ie uademoupensis Post onl34(PC, isotypes tn. Sie® NY).
Wevuadeloupe, M.i;'Herminier". =| Fomitopsis supina (Fr: )
Ryv. as already indicated by Bresadola on the. original
label.

Pemnasvoerl1 i OU 190N ti, isotypes in PCek. BP):
Java. =—sPhellinusspectinatus (K1l.)) Quel. as indicated
on the sheet by Bresadola~

Prenasse titi wl Ouse 187%
The type has not been found (Java).

P. heteromorphus 1846:123.

"Guyana". = Amauroderma schomburgkii (Mont. & Berk.)

Torr., as already indicated by Furtado (1968:268).
erenymenius 1663°263° (PC, isotype in’ kK).

chien Mee I OOROU LI etl co0"*CColombial., = Coriolopsis

brunneo-leuca (Berk.) Ryv.

The type: is unfortunately sterile. I have not seen
the species fertile from the neotropics, and the dextri-
noid reaction found in the skeletal hyphae of many Afri-
can specimens was not seen in Léveillé's specimen. It
may be that two macroscopically similar species are in-
volved. Fertile material from the neotropics may shed
some light on the problem.

Mem nouc na tus OuOMLNORC PG isotypes: in FH & BPI).
BeeruoOtvetyanelid Ghnerry, India orientabist.—= Trichap-—
CuUMme DL OrmissnGrreeun Kile)” RyVes

Be kKickxianus 18482122.

The typeshas not been found (America).

Eereorthalsi i091 844-9190 3h):

"Sumatra, Korthals". = Phellinus senex (Nees & Mont.)
Imaz.

feerlcnicwmicloc lesm (PC. isotype in’ BPL)*

"America, ad truncos". = Coriolopsis polyzona (Pers.)
Ryv.

Pomlenziteus 1654-17>°(L, isotype in PC).

"Zollinger 975 (Sumatra)". = Lenzites acuta Berk.

P. leucomelas 1846:140.

The type has not been found. Invalid name, non P. leuco-
melas Pers. 1825.

eo indivi so3e263 (PC, asotype in’ K)).

"Tepuendama (Colombia) Canoas alt 2600. Sept. 1860".

LBZ

= Trametes cfr. modesta (Fr. ) Ryv. The collection is fer=
tile with spores 6.5-8 x 3-3.5 um. These measurements are
in accordance with those of Fidalgo & Fidalgo (1968:27).
The spores of the paleotropical taxon called T. modesta
and macroscopically identical with that of the neotropics,
are seemingly smaller. However, a richly fertile specimen
has never been found and no sporeprint has been made.

P.. dongipes® 184621249 (PC; isotype, in’ BPL)«
"Guyane francaise". This is an accepted species and-the
type of Haddowia, Stey. in the Ganodermataceae. For a
description, see Ryvarden & Johansen (1980:93) with fur-
ther references.

P. macropus 1848:122.
The type has not been found (Java).

P. mangiferae 1846:130 (PC).
"Voayge de M. Perville. N. East de Mahe (Tahiti) 1841".
The species belongs in Ganoderma and its final status
has to be decided when the genus has been revised.

Premanubprelatus foots lyauUler
"Zollinger 974" (Sumatra). = Microporus scopulosus
(Berk.) Ryv.

Pema ren onc sme) od Os 00n
The type has not been found (Marguesas Islands).

Paminastoporus Lolobel ;omure)..
"Singapore, Février 1832". This is a Ganoderma species
im thesG. =huctdum= complex. =Ltsestacvus nas stom bercetered
when the genus is revised.

P. megaloma 1846:128.
The type has not been found (New York, USA). In American
literature it is commonly assumed that this is the same
as Ganoderma applanatum, see Overholts (1953:99). From -
ther déeseription itsistvery probabil yeso-.

P. melanevis1046.131) (linen sotypes dneoes eBrion.
"Zollinger no. 2085 (Java)". = Coriolopsis caperata
(Berk.) Murr.

P. melanaleucus 1846:141.
The type has not been found (Réunion).

Pe imicrocyclius 1049 +168 “CLs wsotypes ine POsiKeeeNY. mb bie)e

"Java". = Cyclomyces tabcinus (Mont.) Pat.

Peomicrolona aaa; 183.
The type has not been found (Manilla, The Phillipines).
At Kew there is a specimen: "Phillipines 1908, Curran,
cum. typo comparato’™ ink, Bresadola's hand. This isva
specimen of Microporus affinis (Nees & Blume ex Fr.)
Kunt. The specimen mentioned above is selected as neo-
typemuntile authenticemateriadwisefounds,

P. Mmorutzianus) 1 o40s1 301 Cla.
"Zollinger 360 (Java). = Trametes marianna (Pers.) Ryv.

Pi. murdinus! 16444185 (i sasotypes an, POsanderi)s
"Java". = Trametes menziesii (Berk.) Ryv.

P. nephelodes 1846:125.

The type has not been found (Paramaribo, Brazil).

In the Stockholm herbarium there is a specimen: "182 P.
nephelodes Lév. = Polyp. flabelliform. Kl. transiens Ja-
va v. Héhnel" Bresadola's hand. The specimen is Micro-
porus affinis (Blume et Nees ex Fr.) Kunt. However, as
the genus Microporus is not known from South America,

b8sS

the specimen from Java cannot be selected as neotype of

P. nephelodes.
P. nordmannii 1842:93.

The type has not been found (Korbek, USSR). Fries
(1874:581) refers the species to Heterobasidion annosum
(Fr...) Bref. but it is not stated whether he had seen the
type or based his: suggested synonymy on the descrip-
tion alone. The latter is rather vague, but could apply
tOVasresupinate specimen ofvH. annosum.

P. notopus 1844:194 (PC).

"Java". = Trametes modesta (Fr.) Ryv.
Pemostreabusmilodo lo 7m PC erisotype IneBel).

"Bresil, Rio de Janeiro, M. Gaudichaud 1831-33". = Tra-
metes cubensis (Mont.) Sacc.

P. pala 1844:183.
The type has not been found (Surinam).

Peepeetunculus! O46413 acre).
"Indes orientalis, Nelli-Gherry (India)". = Trametes
versicolor (Fr.) Pil.
P. perpusillus 1844:191 (PC).
"Am. Boreal". = Fomitopsis scutellata (Schw.) Bond. &
Sing. ar
P. phaeus 1846:132.
The type has not been found (Sri Lanka).
Peeplacopusraldtbalotas )i
"Java, Junghuhn". This is a Ganoderma sp. in the Gano-
derma lucidum complex. Its status has to be settled when
the genus is revised.
P. platypilus 1844:192.
The type has not been located (Java).
Pee pkumbeus: 10462136 (PC, Asotype inwS).
"Guadeloupe, M. L'Herminier". = Rigidoporus fusco-linea-
CUSMULECS ony VE
P. rhodophaeus 1844:190 (S).
"Java". This is an accepted species in Fomitopsis as
F. rhodophaeus (Lév.) Imaz. For a description, see Ry-
varden & Johansen (1980: 340).
P. rigidus 1844:189 (L).
"Java, Zollinger no. 732". = Nigroporus durus (Jungh.)
Ryv.
Rea Pudis a. C4643 30 (PC).
"Guadeloupe, M. L'Herminier". Invalid name, non P. rudis
Berk. 1839. The type of the invalid name is Fomitopsis
BUDS BOE eh VV.
P. rugulosus 1844:189 (L).
NNow57'" CJavaja(leg. Junshubn?,). i= Coriolopsis sangui—
naria (Kl.) Teng.
P. seclerodermus 1846:129 (PC).
"Guadeloupe, L'Herminier". = Fomes fasciatus (Fr.) Cooke.
P. ‘scleropodius .1846.123 .(BPI.).
"Tle Bourbon (Réunion)". = Lignosus sacer (Fr.) Ryv.
P. sericellus 1846:125.
The type has not been located (Sri Lanka).
Pawetoeroides 104u lee (PCeisotype in kt).
"Sumatra". This is an accepted species in Coltricia as
G. sideroides (Lév.) Teng., for a description, see Ry-
varden & Johansen (1980:112). Léveillé named also Coll.

184

Zollinger No. 90 as P. sideroides, but this collection
is a specimen of Phylloporia chrysita (Berk.) Ryv. Dup-
licates of this collection have been seen in herb. L,
Seer rigger On and mb inks
P.-sordidus- 16442192 (PC) isotype ins).
"Guadeloupe, L'Herminier". = Trametes modesta (Fr.) Ryv.
P. splendens 1844:187 (PC, isotype in L).
"Java". = Trametes modesta (Fr.) Ryv.
P. spurcus ,1946°1350(PCy isotypesvin’ S)-
"Guadeloupe, L'Herminier". = Phellinus gilvus (Schw.)
Parts
P. stevenii 1842:91.
The type has not been seen. Lloyd (1915:285) indicates
it to be Ganoderma applanatum based on the original des-
cription and plate. Donk (1974:67) agreed, and I sub-
scribe to the conclusion.
Pe subf lavusau c4or 300m (PC)E
"Nouvelle Orleans, Salle". = T. versicolor (Fr.) Pil.
P. swartzianus 1846:132.
The type has not been found (Jamaica).
Po tegularise1 040-1 319eGPC mn SOLy peal tmnt).
"Zollinger 2054 (Java)". = Trametes scabrosa (Pers.)
Cunningh.
Piemtvenaxelo40 >: 139%
The type has not been located (Teguendama, Colombia).
P.. tener 16465139 CPC)e
"Guadeloupe, M. L'Herminier". = Trametes villosa (Hook.)
Ryv.
P. tenuissimus 1844:188.
The type has not been found (Java, leg. Korthals).
Petestaceus 91 S46" 126g(PC)™
"Brasilia meridionalis, Dupré, 1842". This is a Ganoder-
ma sp. in the G. lucidum complex. Its status has to be
settled when the genus has been revised.
P. trachodes 1844:192.
The type has not been found (Java).
Pemetbricolor slot be 34e
The type has not been found (Bahia, Brazil).
Petunia conus 640-15 deer cee
"Zollinger No. 2069 (Java)". = Coriolopsis sanguinaria
(Kis mere np.
Pamitbris ti smlo4b tc OmGRpCeaeisot ypesinakhy
wZoLVnger |\No-=2035° (ava )feuinvaliduname,, nontr.ntris—
tis Pers. 1825. The type of Léveillé's invalid species
is a specimen of Nigroporus vinosus (Berk.) Murr.
P. unguiformis 1846: 13% (PON@isotypemain fH).
"Zollinger No. 1437, Tjiboya (Java)". = Rigidoporus mi-
croporus (Fr.) Overeem.
Peoevolneratusml cn veloc ec.
"Ad truncos, Java". = Coriolopsis sanguinaria (K1.)
Teng.
P. zollingerianus 1846:131 (PC, isotypes in S and FH).
"Zollinger 1386 (Java)". = Trichaptum byssogenus
(Jungh.) Ryv.

185

SISTOTREMA Fr.

S. ochroleucum 1846:145 (PC).
"Bombay (India), Polydore Roux" = Lenzites acuta Berk.

TRAMETES Fr.

ieeacuta 1LoW4s196.
The type has not been located (Sumatra). Specimens deter-
mineduby sbréesadolavin herb. So and Bri eare all Coriolop—
sis strumosa (Fr.) Ryv., and this was probably also Lé-
veillés concept.

Tercrassanouy soya ko).

"Madagascar". = Hexagonia hirta (Fr.) Fr.

T. dermatodes 1844:196 (PC).
"Voyage de M. Gaudichaud sur la Bointé. 1836-37. Manille
(The Phillipines)". = Oxyporus cervino-gilvus (Jungh. )
Ryv. as already indicated by Reid. (1953:135).

Pei ncanagi cud] 06m Bre
"Manille (The Phillipines)". This is an accepted species
in Trametes, for a description, see Ryvarden & Johansen
(1980:565).

ieeperrottetiiy 16441 95 (PC).
"Guyane francaise M. Poileau" (printed on the label).
"Java, M. Perrottet", unknown handwriting. The name,
however, is written by Léveillé.

There has been a mixing of labels or facts. The type
represents a South American taxon which has never been
found or reported from the paleotropics. The type is ap-
parently the same specimen which was used by Montagne
and Berkeley when they described Trametes trichomallus
in 1849. The latter is the type of Trichaptum Murr.

Trichaptum perrottetii (Lév.) Ryv.
NOOW sted ee DOL elo 23 (an Lope.

Fruitbody applanate, sessile, semicircular to elongated
shelf-like, mostly broadly-attached, usually not decur-
rent on the substratum, 5-15 cm long, 3-/ cm wide and

to 8 mm thick (tomentum not measured), tough and flex-
ible. Pileus with a dense, strigose to villose or hir-
sute layer of forked hairs, dark brown, becoming darker
towards the base and more greyish towards the margin,
azonate or weakly-zonate, up to 10 mm thick at the base.
Margin entire and sharp. Pore surface snuff brown, pores
angular to round, first entire and thin-walled, 2-3 per
mm, in older specimens often incised, coalescing and in
parts sinuous to daedaleoid, in the latter case up to 2
mm wide and several mm long. Tubes deep brown, 2-5 mm
deep. Context very thin, 0.1-0.4 mm, brown to dark och-
raceous. Hyphal system dimitic, generative hyphae thin-
walled, hyaline and with clamps, 2-4 um wide. Skeletal
hyphae abundant, thick-walled to solid, mostly yellowish
to light brown, 3-5 um wide. Cystidia abundant in the
hymenium, clavate to ventricose with a tapering apex,
smooth or with an apical crown of crystals, 10-18 um
long, slightly projecting, basidia clavate 12-15 x 4-6
um and with 4 sterigmata. Spores subcylindrical to ob-

long ellipsoid, smooth, thin-walled and non-amyloid,
5-7 x 2-3(3.5) um. On deciduous wood.

Specimens have been examined from USA, Cuba, British
Honduras, Guatemala, Costa Rica, Colombia, Brazil,
French Guyana and Bolivia.

Tee vel tba var hod delOG™.
The type has not been found (Sumatra).

ACKNOWLEDGMENTS

The staff of the herbaria in Stockholm, Leiden, Paris,
Kew, New York, Harvard and Beltsville are most kindly
thanked for their cooperation during my type studies. The
Norwegian agency for International Development has given fi-
nancial support for which I am very grateful. R. Winter has
suggested improvements in the English text.

REFERENCES

Bursdalil eH. Heir. SaMiler.) Oserk.n OO rl ereevaluuatdormon
Panellus and Dictyopanus. Beiheft. Nova Hedw. 51:79-91.

Donk, M. A. 1974: Check list of European polypores. Verhand.
Afd. Naturkund. Kon. Nederl. Akad. Wetensch. 2 ser. no 62.

Pidalgo, eM. ies Pak. kwridal goles 0.01960 sePokvuporacescmiicom
Venezuela I. Mem. New York Bot. Gard. 17, no 2:1-34.

Fries, E. 1874: Hymenomycetes Europaei. Uppsala.

Furtado, J. S. 1968: Revisao do genero Amauroderma (Polypo-
raceae). Sao Paulo. Priv. ed.

Léveillé, J. H. 1842: Observations medicale et enumeration
des plantes, p. 35-237 in Demidoff, A. (ed.) Voyage
dans la Russie meridionale et la Crime. Vol. 2.

Léveillé, J. H. 1844: Champtgnons exotigques._ Ann. Sei. ‘Nat.
Det een. lem eO Giclee.

Léveillé; J. H. 1846: Description des champignons de l'her-
bier vdu Museumsede) Parts Ann.) ooCl Na Camel es my Oem
ea oy a

Léveillé, J. H. 1846b: Fungi, p. 164-204 in Gaudichaud, M.
(ed.) Voyage autour du Monde sur la corvette La Bonité.

Botanidue.9 V0. ous lk.

Léveillé, J. H. 1848: Fragments mycologiges. Ann. Sci. Nat.
Ser. 3. Vols. '9:119-144.

LEevetie gc) eh. sLood ce hung igeip. Bobo ZOlsme riper mtd s)
Systematische Verzeichnis der im indischen Archipel in
Jabren 1842-48 gesammelter sowie der aus Japan emp-
fangenen Pflanzen.

Léeves ese J eH. e166 3 ceehunet Sine teriang eM. wnieues eel anchors
J. E. (ed.) Florae nova-granatensis. Ann. Sci. Nat.
Sere UmVol. 20262-3800)

Lloyd, C. G 1912: The polyporoid types of Léveillé at Lei-
den. Meded. Rijks Herb. Leiden 9:1-5.

Lloyd, C. G. 1915: Synopsis of the genus Fomes. Lloyd Mycol.
Writ. 24 eo ld —206.

Reid, D. 1953: New or interesting records of Australasian
Basidiomycetes III. Kew Bull. 12:127-143.

Singer, R. 1945: The Laschia-complex (Basidiomycetes).
Lloydia 8:170-230.

MYCOTAXON

VOT AL ELE Nomal ppl 87-190 April-June 1981

THE PUBLICATION DATE OF ARENDHOLZ'S THESIS
ON LEAF-INHABITING HELOTIALES

WOLF-RUDIGER ARENDHOLZ

Biologie, Universitat Kaiserslautern
6750 Kaiserslautern, West Germany

and
RICHARD P. KORF

Plant Pathology Herbarium, Cornell University
ithacay. New!) York 14853 7USA

SUMMARY

July 7, 1979 is shown to be the date of effective pub-
lication of W.-R. Arendholz's doctorate thesis, 'Mor-
phologisch-taxonomische Untersuchungen an blattbewohn-
enden Ascomyceten aus der Ordnung der Helotiales," and
the date of valid publication of the new taxa and new
combinations proposed therein.

Tiveesta usmiote pUDlICationj#o!l BWs—Rk ae Arendholz!s | Pheb:
thesis, entitled "Morphologisch-taxonomische Untersuch-
ungen an blattbewohnenden Ascomyceten aus der Ord-
nung der Helotiales'' was assumed to be "unpublished"
Pree KOnimalcdenkOnneulLjooymemmmorder tOmprotectthesstatus
Of VArendholz.s. new names, they reported) one, as an
as yet unpublished new species referred’ to the genus

~w slhey ‘shad received in’ the mails from’ Arendholz
a copy of his thesis (Arendholz, 1979), a soft-bound
volume having the appearance of being some sort of
photo-copy of the original typewritten thesis on deposit
with the Universitat Hamburg.

The purpose of this note is to call attention to the fact
that over 180 copies of this thesis were actually distri-
buted. eandy inateiny no, way can, the» new, names, therein
be ignored or treated as not validly published. On July
7, 1979, 139 copies were mailed by Arendholz to the

188

Staats— und Universitdtsbibliothek, Hamburg, which sub-
sequently distributed them to other libraries, and that
date is held here, to constitute the (date otmdistri bution
CONSLITUTINGeellectivempublications sundeissticasbnicomdil
onal Code of Botanical Nomenclature (Stafleu et al.,
LOU Seem Gt Cc lome2 J .61% Twenty further copies were sent
tO the = Wibranyemol mache? SINStitutiethuL ee |CemelneeeDOLduik
ders Universitat Hamburg ss andysanother a 2 5mund cee il) samec
scientists in different countries. Approximately) 20 addi—
tional @ copies: pancees Ea nvweALendn Olzcm DOS SCSSoIOUmm OULU
of a photo-offset lithography press run of some 200
cOpLes.

While it seems unfortunate. that.the copies that were dis-—
tributed) Camry, ino indications of sthesexactsddatesOlmars ume
Dution, —nor Vveven wthats themvolume sconsuitMies samspubDlica—
tion in the sense of the Code of Nomenclature, recent
articles on “effective” publication »under VArticles® 29-3 lmot
the ~Codestleavem nowdouDl ming our nincde ta teethometiesis
was jcifectively, published men! Nemes fOurematliCleSmal imal acon
29(4), the August 1980 issue, support our _ position
[Brummit (1980); Hara and Eichler (1980); Nicholson
(1980); Weresub and McNeil (1980) ].

Since thessthesisieiS = printed byes pnoto-Ofisetmlithograpiye
it must apparently be accepted as printed matter (Art.
29): “... it may safely be assumed that matter pro-
duced’ *' by 9a conventional Gtypewriter, « Consisting Olgsone
Lope copy, plusmone. to several carbon scoples. =i sanetncol.
sidered sto) be, printed. On ties other ihand = copyeos ginal
ly produced on a typewriter and then reproduced by off-
set lithography (as.for example the important taxonomic
journal Mitteilungen der Botanischen Staatssammlung
Miinchen) must certainly be accepted as printed matter"

(Brummitts 1980-87 =4/0)8

The, Codes (Art 9291) requires sthateithes printeds matter
bes@diptributedmtomthemoceneralmpublicwoumdteslcastmtompo—
tanical institutions with libraries accessible to botan-

iStSeeoercna | Ly. eNed (Liem tiem Un DelmOtmcOD1es mmo tmL Tc
number of libraries: (nor even that they be located in
more than one country) is stipulated in the Code, "and

presumably in an extreme case a printed work distribu-
ted to as few as two libraries should be taken as ef-
fectively published" (Brummitt, 1980: 479).

In the case in point, the requirements for effective pub-
lication would have been fulfilled by mere deposition
of one copy of the printed thesis in the "Staats— und
Universitdtbibliothek" and one in the library of the '"In-

189

sxiiuteeitir “Allgemeine ~—Botanik.” As noted above, many
more than two copies were distributed to several libra-
ries and scientists worldwide. Nicholson's (1980). key
Similarly keys out this thesis as "effectively published."

Establishing an exact date of publication of any prin-
heaesworkt is icritical for nomenclatural purposes. For
journals, the practice is to use’ the actual date of mail-
ing of each issue as publication date [this is the
procedure with MYCOTAXON, as it is also with MYCOLO-
GIA (C.T. Rogerson, Managing Editor, pers. comm.) ].
Similarly, commercial books are published on the date
they are actually offered and available for purchase,
not that on which they actually reach the public or li-
braries" (Brummitt, 1989: 480). Since the bulk of the
copies of Arendholz's printed thesis were mailed on
Puy on? je) 19/9, tom the Staats—) und Universitatsbibliothek,
HagiDunewathats date shouldmnow be “accepted as the date
of publication of the thesis for all nomenclatural pur-
poses.

Various proposals by Hara and Eichler (1980) and by
McNeil (in Weresub and McNeil, 1980) focus on the kind
Ol problem presented by ‘Arendholtz’s thesis with a view
tO eliminating any problems in the future, with worth-—
while suggestions for requiring an indication of inten-
Gion to molfer material “as ~“*published’” under the provi-—
PiencmoOtmiies COde,marequirements {fonsdeposition im ajspe—
cific number of (named) institutions, and indication of
actual date of publication (distribution). Such propo-
sals will be useful in the future, but in any case will
not be retroactive, but apply from 1982 onward if they
are adopted at the International Botanical Congress in
sydney later this year in the forms submitted.

REFERENCES CITED

ARENDHOLZ, W.-R. 1979. Morphologisch-taxonomische Unter-
suchungen an blattbewohnenden Ascomyceten aus der Ord-
nung Helotiales. Ph.D. Thesis, Univ. Hamburg, 115 pp.,
fom mCP ri vatelvyedistributed 7 July 19/79)

BRUMMITT, R.K. 1980. Questions on effective publication.
Taxon 29: 477-483.

HARA, H. & H. EICHLER. 1980. Proposal (67) to amend Art.
29.1 on effective publication and recommendation to esta-
blish a special committee on Art. 29. Taxon 29: 515-516.

MOR AER Pec lte MM KOHN peu lO7G |) Revisionary studies “inethe
Hyaloscypheae. I. On genera with "glassy" hairs. Myco-
taxon 10: 503-512.

190

NICHOLSON, D.H. 1980. Key to identification of effective-
ly/ineffectively published material. Taxon 29: 485-488.

STAFLEUs, F-As. et al. feds.|] 1978. Internationakecodesot
botanical nomenclature adopted by the twelfth Interna-
tional Botanical Congress, Leningrad, July 1975. Regnum
Veg. 197 :9i-xiv, 1-457.

WERESUBY OLN Ke & 0 Hea McNeilo) 1980.) Et recuives publicatron
under the Code of Botanical Nomenclature. Taxon 29: 471-
476.

MYCOTAXON

VOLT OMULE NO we ppeato1-240 April-June 1981

AN ALTERNATIVE VIEW OF CERTAIN
TAXONOMIC CRITERIA USED IN THE
ENTOMOPHTHORALES (ZYGOMYCETES )

RICHARD A. HUMBER

USDA-SEA-AR Insect Pathology Research Untt
Boyce Thompson Instttute
Tower Road, Ithaca, New York 14853

SUMMARY

Long-standing unresolved controversies about the
various generic classifications of the entomopathogenic
Entomophthorales require a review of the validity and the
relative weights of the morphological and other taxonomic
criteria used to construct a useful classification for this
group. Major morphological criteria used in the Batko and
Remaudiére classifications are evaluated here in a survey
of the taxonomic distributions of these characters and the
Significance of the irregularities in their distributions.

The number and nature of nuclei in the primary spores,
the branching of sporophores, and the mode of discharge of
primary spores are regarded as the major characters suit-
able for delimiting genera. The presence of rhizoids and
(to a limited extent) the shapes of primary spores are
Significant secondary generic characters. The presence of
cystidia, mode of resting spore formation, and presence of
capillary secondary sporophores are not regarded as char-
acters of generic importance. The conidial or sporangial
nature of primary spores is discussed, as are difficulties
in the interpretation and applications of the terms 2ygo-
spore and azygospore.

bo2

The applications of the generic names used in the
Batko and Remaudiére classifications are reviewed in light
of the findings reported here. The validity of both
Stronguellsea and Entomophaga is upheld, and the sense of
Erynia is restricted to those species having branched
sporophores and uninucleate bitunicate primary spores which
do not form secondary spores on capillary sporophores. A
nomenclatural problem is noted which indicates the synonymy
of Zoophthora and Erynta if these genera are not retained
as separate as proposed in the Remaudiére classification.
Tabanomyces is regarded as a synonym of the nematophagous
genus Mertstacrum.

INTRODUCTION

A far-reaching revision of the entomopathogenic genera
of the Entomophthorales (Zygomycetes) proposed by Batko
(1964a-e, 1966; Batko and Weiser 1965) has spawned consid-
erable controversy among the students of these fungi. Even
though there is now wide agreement that it is unsatisfac-
tory to continue classifying approximately 150 diverse
species in the single genus Entomophthora Fresenius, there
has been no agreement regarding the appropriateness of the
criteria used by Batko to restrict Entomophthora and to
establish segregate genera or of the circumscriptions of
these genera.

King and Humber (1981) offer a basic reconsideration
of the major morphological characters of this group and
discuss their taxonomic significance. While King and Hun-
ber evaluate the basic strengths. and weaknesses of the
Batko classification, they consider that no definitive re-
working of the genera would be appropriate without a final
resolution of the long-standing problem of delimiting
Contdtobolus Brefeld from Entomophthora sensu lato.

More recently, however, a major reworking of Batko's
generic classification by Remaudiére and Hennebert (1980)
and Remaudiére and Keller (1980) — hereafter referred to
as the Remaudiére classification — resulted in generic
assignments for some species which led to no manifest in-
crease in the biological homogeneity of the recognized
genera. Many taxonomic decisions in the Remaudiére classi-
fication, and particularly those regarding the validity and

Eos

importance of various characters used to construct classi-
fications of these fungi, were presented without explana-
mroueend, in tury, have triggered further controversy and
alternate views on classification (Ben-Ze'ev and Kenneth
1981a; Humber 198la).

The time is appropriate to examine thoroughly those
morphological and developmental characters which have been
employed in the modern classifications proposed by Batko
and by Remaudiére and his colleagues. The values of these
criteria are examined in the contexts of the available
classification systems. Indeed, a sense of the relative
values of their chosen criteria does emerge from a consi-
deration of the major schemes of entomophthoralean genera
and of the internal inconsistencies generated by the selec-
tion and ranking of the criteria accepted as valid in each
of these generic schemes.

Despite 1ts undisputed desirability, no general agree-
ment regarding the generic classification of the Entomoph-
thorales will be possible until the morphological and other
emicreria used ini the past or being proposed now have been
critically reviewed, evaluated, and agreed upon. This
paper is the first in a series discussing the merits and
weaknesses of the Batko, Remaudiére, and other emergent
Slassitications: (€.g., Ben-Ze'év and Kenneth 198la) in an
attempt to find rational bases on which to build a generic
scheme which (1) avoids the inconsistencies of the previous
classifications of the Entomophthorales, (2) can accurately
predict the natures of characters or aspects of the host-
pathogen relationship, and (3) reflects what appear to be
the evolutionary relationships among these fungi.

TAXONOMIC CRITERIA: ANALYSIS AND CONCLUSIONS
PRIMARY SPORES
Primary Spores: Conidia or Sporangia?

Throughout this paper I refer to primary spores or
spores rather than to the more conventional contdium or the
less common sporangium or sporangtole. Contdtwn and spor-
angtum are terms referring to specific developmental pat-
terns; their use should be for purposes of accuracy and
must be backed by sufficient demonstration of the mode of

194

sporogenesis. The reference to any asexual propagules in
the Zygomycetes as contdta has been questioned by the stu-
dents of the Hyphomycetes and Coelomycetes, although there
is no reason to believe a prtort that conidia cannot be
formed by these or any other lower fungi. If appropriate
studies of the origins and fates of the various wall layers
of spores (and sporophores) during sporogenesis and germi-
nation prove that,indeed, they are conidia (or sporangia),
then these propagules should be referred to as such. Simi-
lar studies are also needed for the panoply of secondary
spore types produced in the Entomophthorales.

Spores containing single nuclei and borne on branched
(or simple, in Strongwellsea Batko §& Weiser emend. Humber)
sporophores have caused much interpretational difficulty.
These spores have a layer which, in liquid, lifts away from
the spore surface and balloons outward. This has been re-
garded variously as a gelatinous coat (é.g., Entomophthora
gloeospora Vuillemin 1886) or, more frequently, as evidence
for the sporangial nature of these fungi by regarding the
detached layer as a sporangial wall enclosing a single
Sporangiospore (Thaxter 1888; Strong et al. 1960; Batko
1964b,, 1974)... This typéesot primarysspore seems, Lompcscn=
tirely restricted to the species of Zoophthora Batko,
Erynta Nowakowski, and Strongwellsea. Kenneth (1977)
claims that the outer wall of Entomophthora turbinata Ken-
neth [= Weozygttes turbtnata (Kenn.) Rem. §& Keller, a gen-
eric assignment which remains debatable] lifts away but
admits (personal communication) that this feature is doubt-
ful and requires further investigation. If the spore of
E. turbtnata truly were bitunicate (with a separable outer
wall layer), it would be the only instance of this charac-
ter in a Species with multinucleate spores.

Humber (1975; King and Humber 1981; Roberts and Humber
1981) notess ultrastructuralyevidences that thessporesos
Strongwellsea magna Humber, which has a separable outer
wall layer, cannot be a monosporic sporangiole. The spore
wall has no indigenous (sporangiospore) layer that is not
shared with) thes sporophorepitse lt ainesseparab les lavenmars
not a distinct (sporangial) wall, but represents only a
separation of the outermost layer of the sporophore wall,
all layers of which are continuous over the entire spore
surface except for the papilla — which is covered only by
an extension of the inner wall layer of the spore and
sporophore (Humber 1975). This spore might be a true coni-
dium, but this cannot be confirmed without also determining

LoS

the fates of the wall layers during germination (see
Dykstra 1974).

The only evidence for the occurence of sporangia in
the Entomophthorales comes from several species of Bastdto-
bolus Eidam, in which there are numerous instances when the
cytoplasm inside a cell (usually but not always a spore)
may be divided into several independent uninucleate units;
B. mtcrosporus Benjamin (1962) provides the most striking
examples of this behavior.

I use the terms unttuntcate and bttuntcate introduced
by Remaudiére and Hennebert (1980) and Remaudiére and Kel-
ler (1980) to refer to spores on which a separable outer
woeelayer siseei ther absent, or present, respectively:

These terms are used almost exclusively within mycology to
refer to the morphology (and function) of asci in the Asco-
mycotina, and constitute a primary character dividing the
(bitunicate) Loculoascomycetes from all other (unitunicate)
classes of this subdivision (Ainsworth et al. 1973). The
use of these terms, however, should be generally accepted
for the Entomophthorales since they (1) are convenient, (2)
respect their etymological derivations, (3) should not be
in any way confused with their applications for the asco-
mycetes, and (4) in the case of bitunicate spores, would
help suppress the notion that such spores are sporangial

in nature.

Morphology

The variety of shapes of primary spores in the Ento-
mophthorales has been the source of much attention, parti-
cularly among those who have included nearly all entomo-
pathogens of this group in the single genus Entomophthora.
A series of artificial species groupings based on spore
morphology have been proposed primarily to aid species
identification (Lakon 1919; Hutchison 1963; Gustafsson
1965; MacLeod and Muller-Kogler 1970, 1973; MacLeod et al.
1976; Waterhouse 1975; Zimmermann 1978). None of these
authors, however, intended their groupings to have taxono-
mic value. Remaudiére and Keller (1980), however, chose
the morphology of primary spores to be the main criterion
for their generic classification.

Several genera do have characters of spore morphology
which are unique to themsleves. One of the strongest and
most valid criticisms of the Batko classification, in fact,

196

was its separation of the species having campanulate spores
(with relatively flat bases and a strongly apiculate apex)
into Entomophthora sensu stricto and Cultetcola Nieuwland
depending upon the absence or presence of rhizoids. Remau-
diére and Keller (1980) rejected the significance of rhi-
zoids for defining the genera, and redefined Entomophthora
to include only species with this distinctive campanulate
spore shape and associated mode of spore discharge (see
MacLeod et al. 1976; Samson et al. 1979).

The spores of Triplosportum (Thax.) Batko (nom. gen.
conserv. prop., see Humber et al. 1981; = Weozygttes Wit-
laczil) are variable in shape, but most often are round to
slightly elongate and have a truncate rather than a conical
or rounded papilla. As circumscribed by Batko (1964c), the
spores of Cultctcola species could be either campanulate
(like those of #. muscae (Cohn) Fres.) or obpyriform with
a smooth apex and rounded papilla; species with the former
type were restored to Entomophthora s.str. while those with
the latter type of spores were transferred to Contdtobolus
by Remaudiére and Keller (1980). King (1976b, 1977)
regards Contdtobolus species to have round to pyriform pri-
mary spores, and notes the similarity of several entomo-
pathogenic Entomophthora (sensu lato) species with Contdto-
bolus. Remaudiére and Keller (1980) extended King's con-
cept of Contdtobolus by transferring all entomopathogenic
Species with rounded to pyriform spores to that genus. The
species of Entomophaga Batko have pyriform spores with a
rounded apex and closely resemble the spores of Contdto-
bolus species. These two genera are unambiguously differ-
entiated, however, by both the morphology and stain reac-
tions of their nuclei (Humber 1981b).

The primary spores of Zoophthora Batko sensu Remaudi-
ére and Hennebert (1980) (= Zoophthora subg. Zoophthora
Batko sensu Ben-Ze'ev and Kenneth 198la) are rather cylin-
drical and usually taper apically to a blunt cone or a
poorly defined point; just above the conical papilla, the
spore flares to a very slight (but not always apparent)
shoulder. The spores of Erynta Nowakowski species (see
Remaudiére and Hennebert 1980) display the greatest shape
variation of any genus in the Remaudiére classification.
They range from elongate fusiform with a marked curvature
and a shallow rounded papilla to somewhat cylindrical to
ovoid or fusiform spores with rounded apices and papillae
canted away from the spore axis. The spores of Strong-
wellsea species fit well within the range of morphological

197

variation found in Erynta, a fact which seems to have moti-
vated Remaudiére and Keller (1980) to merge this genus into
Erynta. The spores of Empusa carolintana Thaxter, which
Remaudiére and Hennebert (1980) transferred to Frynta, are
obovoid to pyriform, a shape corresponding more with those
of Hrynta spores than with any other genus.

Karyology: Number and Type of Nuclei

Early studies noting the taxonomic importance of
entomophthoralean nuclei have been largely ignored (Vuille-
min 1895; Cavara 1899a-b), so that no surpassing importance
was attached to the number of nuclei in primary spores
until the proposition of Batko's classification (1964a-d,
1966; Batko and Weiser 1965), which necessarily drew upon
the observations of earlier workers regarding this cyto-
logical detail.

One of the most significant characters used by Batko
(1964b) to establish Zoophthora was that of uninucleate
primary spores, a character which is readily detectable and
seems to be correlated well with the other characters of
this genus. Strongwuellsea Batko §& Weiser (1965) was also
described as having uninucleate spores but was clearly dis-
tinguished from Zoophthora by its simple sporophores and
unique habit. EHrynta Nowak. was split from Zoophthora by
Remaudiére and Hennebert (1980) and is circumscribed so as
to allow multinucleate primary spores even though £. caro-
lintana is the only such species included in E£rynta by
Remaudiére and his colleagues. As discussed at length be-
Low, thesmorphological icharacters of this species do not
appear to belong in this genus, and its probable misclassi-
fication is clearly indicated by its status as the only
species with multinucleate spores in this large grouping
of species with uninucleate spores.

As is the case with branched sporophores (see below),
there is a very low but natural incidence of abnormally
developed primary spores. The spores of Zoophthora, Erynta,
and Strongqwellsea receive very nearly the entire cyto-
plasmic contents of the terminal cell of the sporophore.

In species with branched sporophores there are, occasional-
ly, larger than usual volumes of cytoplasm containing two
(or three) nuclei which are isolated in a branch by its
basal septum. The spore subsequently formed from this
branch will be both markedly larger than normal and contain
the-extra nucleus’ or nuclei. Such exceptionally rare

L198

spores are clearly aberrant and cannot be used to invali-
date the significance of uninucleate spores. Similarly,
they do not justify the inclusion in Erynta (all of whose
other species have uninucleate primary spores ) of #. caro-
lintana (whose spores are always multinucleate and whose
other major morphological characters do not match with the
other species of Erynta).

Three genera are noted by Batko (1964a-b, 1974) to
have plurinucleate spores. The spores of several Entomoph-
thora species (sensu Remaudiére and Keller 1980) contain
4-6 nuclei; those of £. eultets (Braun) Fres. are charac-
teristically binucleate (or occasionally trinucleate) while
those of E. muscae (Cohn) Fres. contain 5-8 nuclei. Trtplo-
sportum (= Neozygttes) was described by Batko (1964b) to
have quadrinucleate spores. Of the eight species included
in Neozygttes by Remaudieére and Keller (1980), seven have
4-nucleate spores; Entomophthora turbtnata Kenneth (1977)
has spores with 5-7 or more nuclei. This deviation from
the quadrinucleate condition of all other species in this
genus indicates that close scrutiny must be given to the
generic assignment of this relatively little studied fun-
gus. On first examination, the only marked similarities of
E. turbinata with the other species of Triplosportum are
its simple sporophores and ovoid resting spores with a jet
black ‘epispore. Batko; (1974 )eincorrectly minds catedmsnae
the primary spores of Massospora Peck are binucleate; in
fact, they vary from 1-6 (or more) nuclei, but most common-
ly contain 2-3 nucle (Sopers19/455 193i).

The number of nuclei in spores of Cultetcola species
(as originally described by Batko 1964c-d) varied from
pluri- to multinucleate, but really represent a bimodal
distribution with either a few nuclei or many in the pri-
mary spores. This bimodal variation perfectly parallels
the dispersal of Cultctcola species for other reasons to
either Entomophthora s.str. (plurinucleate) or Contdtobolus
(multinucleate) .

The taxonomic importance of nuclear morphology has not
been appreciated until now even though some authors such as
Batko (1964a-b, 1966; Batko and Weiser 1965) have drawn at-
tention to the staining of nuclei in lactophenol/cotton or
aniline blue or other stains, or to their general size and
appearance, but never realized their porential taxonomic
value. The large nuclei of Bastdtobolus species, for exam-
ple, have been the subjects of numerous cytological studies.

MINS

Humber (1981b) finds that nuclei of Contdiobolus spe-
cies (sensu King 1976b) have relatively small nuclei with a
prominent central nucleolus but no obvious heterochromatin;
these nuclei remain unstained or only very weakly differen-
tiated in lactophenol/aniline blue, aceto-orcein, or a nun-
ber of other nuclear stains. The species of Entomophthora
and most of the entomopathogenic genera have large nuclei
with prominent heterochromatin but no large central nucleo-
lus; these nuclei may be differentiated in lactophenol/ani-
line blue and are strongly and rapidly differentiated in
aceto-orcein and other nuclear stains. This karyological
criterion has proven to be of inestimable value for diag-
nostic work, particularly for distinguishing species of
Contdtobolus from Entomophaga (whose primary spores are
nearly identical in shape) or other genera with readily
differentiated nuclei, and in determining incidences of
double infections in insects when one of the fungi is a
species of Contdiobolus.

Taxonomic Significance of Primary Spores

Batko's recognition of the generic significance of the
number of nuclei (one, a few, or many) in primary spores
has been widely embraced. The Remaudiére classification,
however, necessarily disregarded this character in order to
Poace thesgreatestygeneric value on the shapes (of primary,
spores. As will be made more apparent below, this depen-
dence upon spore shapes is misplaced and leads to what seem
to be several misclassifications which offer no improvement
over earlier artificial schemes based on spore shapes and
proposed without taxonomic or phylogenetic significance.

Nuclear cytology affords the only consistent and read-
ily observed criteria which adequately distinguish the spe-
cies of Contdiobolus from Entomophthora and its segregates
(Humber 1981b). These criteria also uphold the validity of
Entomophaga Batko for FE. grylit (Fres.) Batko and several
other species. Entomophthora obscura Hall §& Dunn and £.
thaxtertana (Petch) Hall §& Bell were assigned to Entomo-
phaga (Batko 1964d) but are now regarded as synonyms (see
below); nuclear cytology confirms the transfer of Ento-
mophthora obscura to Contdiobolus by Remaudiére and Keller
(1980).

The unitunicate or bitunicate nature of primary spores
appears to be perfectly correlated with the nuclear number.
With the sole unconfirmed exception of EF. turbinata, all

200

known bitunicate spores also contain a single large nucle-
us. Except for two species of Massospora in which uni-
nucleate primary spores can occur (Soper 1974), all uni-
tunicate spores are pluri- or multinucleate. It is possi-
ble that this distinction in wall structures of primary
spores may assume a greater significance as more entomoph-
thoralean fungi are found and described.

Among the characters related to primary spores in the
Entomophthorales, then, generic value should be accorded to
ae sleast the number and’ nature of the Nuclersitisthesspore:
The shape and uni- or bitunicate nature of the primary
spore should always be noted. However, while various
states of these characters may coincide perfectly with the
generic limits defined by other criteria, no reason is ap-
parent at this time why either of them should be accorded
generic value equal to nuclear number and morphology.

PRIMARY SPOROPHORES

Morphology and Branching

The branching of sporophores was one of the first cri-
teria used to divide the entomopathogenic Entomophthorales
into two genera (Brefeld 1873, 1877; Nowakowski 1883), and
has continued to be one of the leading characters used by
Batko (1964a-c, 1974; Batko and Weiser 1965) to construct a
contemporary classification. The historical significance
of sporophore branching as a character with generic impor-
tance demands careful review of both the reasons for and
the effects of its de-emphasis by Remaudiére and Keller
(1980). It is necessary first to examine the systematic
distribution of simple and branched sporophores and then to
evaluate the exceptions to the normal state in cach genus.

For any practical purpose, the sporophores are always
simple in Entomophthora (although in £. cultets several
Sporophores may arise from each hyphal body, thus giving an
impression of branching), Massospora, Tritplosporium, and
Completorta Lohde, a parasite of fern prothallia. Tabano-
myces Couch et al. (1979; from tabanid fly larvae) and
Mertstacrum Drechsler (1940; from nematodes) produce an up-
right, unbranched sporogenous hypha which becomes septate
and produces forcibly discharged spores on lateral (or ter-
minal) papillae. Ballocephala Drechsler (1951; from tardi-

201

grades) also produces an upright, unbranched sporogenous
hypha on which small lateral cells are formed from which
the spores are budded and forcibly discharged. Sporophores
in species of Ancyltstes Pfitzer (Berdan 1938; from desmid
algae) and Bastdtobolus (Drechsler 1964) may occasionally
branch; those of the 27 Contdtobolus species recognized by
King (1976b, 1977) also may, on rare occasion, be bi- or
trifurcate (King and Humber 1981). Among the species
transferred to Contdtobolus by Remaudiére and Keller (1980)
the sporophores of Empusa aptculata Thaxter and EF. major
Thaxter may show some branching on rare occasion (see Thax-
COnmelooG wert Cf 500) Dut alesal Mos UsaiWayS*Sample.s aneTe .1s
no significant branching in any Entomophaga species.

All species of Zoophthora s.str. (Remaudiére and Hen-
nebert 1980; Keller 1980; Ben-Ze'ev and Kenneth 1981la-b)
have digitately branched sporophores with the possible ex-
ception of Z. crassttuntcata Keller (1980) in which the
sporophores appear in histological sections to be mostly
simple.* All but three Erynta species (Remaudiére and Hen-
nebert 1980; Remaudiére and Keller 1980; Kramer 1981) have
digitately branched sporophores.

Within the bounds of the usual biological variation,
it is not surprising that some species with simple sporo-
phores occasionally have a small number of sporophores that
show a low order of branching. Whatever branching occurs
in these genera is usually basal rather than apical as in
Zoophthora and Erynta (where the apical, digitate branching
pattern usually results in the formation of a tightly
interwoven hymenium). The infrequent occurences of
branched sporophores in genera characterized by simple
sporophores or of occasional simple sporophores in genera
characterized by branched sporophores in no way indicates
that this character should be de-emphasized or discarded.

The sporophores of Zoophthora and Erynta (apart from
the three exceptions discussed below) do exhibit a variable
degree of branching. Unbranched sporophores might occa-

* Keller's (1980) characterization of the sporophores of
Z. crassttuntcata as mostly simple seems to be based en-
tirely on histological sections and should, therefore,
be accepted as provisional until the publication of
micrographs of carefully dissected pieces of hymenium
showing whole sporophores of this species.

202

sionally be found on specimens from this large group of
species in which nearly all sporophores are digitately
branched. Similarly, species such as Z. crassttuntcata

may exist in which (digitately?) branched sporophores are
less numerous than simple ones, but the affinities of these
species with Zoophthora or Erynta will always be apparent
from the natures of those branched sporophores, their pri-
mary spores, and other associated characters.

The natural degree of plasticity in the branching of
sporophores in species of Zoophthora and Erynta does not
imply, however, any real probability of finding species
such as Z. radtcans (Bref.) Batko (= Entomophthora sphaero-
sperma Fres.) or Erynta neoaphtdts Rem. & Henneb. (= Ento-
mophthora aphtdis Hoffman sensu Thaxter) which form obvi-
ously digitate sporophores to yield an exceptional specimen
in which a majority (much less all) of the sporophores are
simple. It is even less feasible that a strain of such a
Species exists which produces branched sporophores when in-
fecting most hosts but which forms only simple ones on one
or more other hosts.

The “uniform occurence of digitatessporophnoresain
Zoophthora sensu Remaudiére & Hennebert — including those
(digitately?) branched sporophores of Z. crassttuntcata —
leaves only the two species of Strongwellsea and EF. caroli-
ntana, the three exceptional species placed in Erynta by
Remaudiére and his colleagues, to account for the de-empha-
sis by Remaudiére and Keller (1980) of sporophore branching
aSpageenericrcr!lernion:,.

It is important to examine why Remaudiére and Keller
(1980) supposed that a significant probability exists for
finding branched sporophores in the two Strongwellsea spe-
cies. It is these fungi which seem to form the basis for
their de-emphasis of sporophore branching. The statement
that branched sporophores are "not always present" in
Stronguellsea, their primary justification for their opin-
ion, seems to draw solely upon Strong et al. (1960) who
note that the sporophores are "rarely branched" but who
also admit the difficulty of tracing complete structures in
their prepared slides. Batko and Weiser (1965) used some
of these slides and correctly described the sporophores of
S. castrans Batko §& Weiser (1965) to be unbranched; they
neither illustrated nor noted any exception to this simple
State. Humber (1975, 1976) also found the sporophores of
S. magna Humber to be always simple, and included this

203

characterization in the emended generic description of
Strongquellsea. The uniformly unbranched nature of sporo-
phores in this genus reflects the means by which additional
sporogenous hyphae interpolate themselves into and thus
continually expand the surface area of the fungal ball in
the host fly's abdomen (Humber 1975).

It has already been noted that #. caroltntana (with
multinucleate and unitunicate spores, simple sporophores,
and lack of rhizoids or cystidia) shares no major charac-
teristics with Hrynta species except for spore shape; the
SimnplessporophnoressOtetnisesnecies do notconstitute any
Significant exception to the common state of digitately
branched sporophores in Erynta. However, some minor degree
of branching of £. caroltntana sporophores may occur t1nstde
the host body, but the emergent sporophores are uniformly
unbranched on the surface of affected craneflies (Thaxter
1888). Both Giard (1888, as EF. arrenoectona Giard) and
Keller (1978) also find the sporophores of FE. carolintana to
be simple.

Several aspects of sporophore morphology — the pre-
sence and pattern of branching and the overall shape of the
sporogenous cell or sporphore apex — should be considered

to be important, but generic significance has usually been
placed only on whether a sporophore is simple or branched.
The sporophores of all Entomophthora species (in the strict
sense) are markedly swollen and clavate below the spore, but
this is the only genus in which this character is uniformly
Dresent a nero 1s58a ess consiscent tendency, to produce
clavate sporophores in species of Zoophthora or Erynta. In
most other species and genera of the order, the apical por-
tion of the sporophore is relatively cylindrical with only
a slight tendency to become clavate.

Srinivasan et al. (1964) proposed to separate Contdto-
bolus from Entomophthora on the basis of sporophore shape.
They suggested that Contdtobolus species have micronemous
(hypha-like, indeterminate) sporophores while those of
Entomophthora are macronemous (thicker than and distinctly
differentiated from the vegetative hyphae, and of determi-
nate length). This criterion was devised to separate
Contdtobolus species from Entomophthora muscae (the type of
its genus) and paid little attention to the many other spe-
cies in Entomophthora sensu lato. In the more modern taxo-
nomic systems considered here, the criterion proposed by
Srinivasan et al. (1964) is applicable only to the few spe-

204

cies of Entomophthora s.str. (Remaudiére and Keller 1980);
it is also untenable, however, since Contdtobolus adtaere-
tus Drechsler (see King and Humber 1981, Fig. 4f) and some
other Contdtobolus species have markedly differentiated
(macronemous) sporophores.

Several genera can, in fact, be distinguished in part
by the exact morphology of the sporophore. Bastdtobolus
species have a more or less prominent swelling of the
sporophore immediately below the spore, and the sporophore
is markedly narrowed at the point where the spore is at-
tached. Species of Meristacrum, Tabanomyces, and Ballo-
cephala form several spores on each unbranched erect sporo-
genous hypha. The mode of spore production in Mertstacrum
and Tabanomiyjces 1S identical, and these genera are here
considered to be synonymous (see below). The spores of
Ballocephala are produced on globose to elongate lateral
cells produced sympodially at the growing apex of the
sporogenous hypha (Drechsler 1951; Richardson 1970; Pohlad
and Bernard 1978). Three species placed in Contdtobolus by
Remaudiére and Keller (1980) — &. apteulata, E. major, and
E. paptllata Thaxter — differ from all other species of
that genus {in the sense of King 1976b, 1977) by having an
extended and neck-like narrowing of the sporophore apex;
this morphological difference suggests that ally characters
of these fungi should be compared carefully with those of.
Contdtobolus before accepting their placement in this par-
ticular genus.

Taxonomic Significance o1° Primary vporopnores

The presence of branched or simple sporophores was re-
garded as a primary generic character in the Batko classi-
fication but was de-emphasized (and effectively rejected)
in the Remaudiére classification, seemingly to justify the
synonymy of Strongwellsea (in which the sporophores are al-
ways simple) with Hrynta (in which the usual case is for
sporophores to be digitately branched). This de-emphasis
and the elevation of spore morphology to a primary generic
character by Remaudiére and Keller (1980) further confused
the classification by placing £. caroltntana (with simple
sporophores) in Erynta and Entomophthora carpenttert Giard
(apparently with branched sporophores, see below) in
Contdtobolus.

As has been advocated by Ben-Ze'ev and Kenneth (1981a)
sporophore branching should be re-instituted as a prime

ZOS

generic character. This restoration sweeps aside the ap-
parent misclassifications of the Remaudiére scheme by re-
turning Strongqwellsea to its rightful status as a separate
genus (Humber 1975, 1976, 198la), and leaving the place-
ments of £. caroltntana and EF. carpenttert undetermined
andgineneed Of sturtner study.

For the entomopathogenic genera and Contdtobolus, the
most taxonomically significant aspect of primary sporo-
PaOLVessISeODLyY whetheruchicy are simpte OF Drancned, There
does seem to be a significant difference, however, between
a low level of basal branching (as in Contdtobolus and E.
earolintana) and the apically digitate branching restricted
to the species of Zoophthora and Erynta. The means by
which the primary spore or spores are produced on the
sporophore and the morphology of the sporophore apex also
assume taxonomic significance when considering the complete
spectrum of entomophthoralean genera.

MODE OF FORCIBLE DISCHARGE
OF PRIMARY SPORES

Three distinct mechanisms for the forcible discharge
of primary spores are known from the Entomophthorales. Two
Ouethesc mechanisms appear to be entirely restricted. to
Single clearly defined genera while the predominant mecha-
nism operates in nearly all the remaining genera. Primary
spores are forcibly discharged in all generally recognized
meonera Of this order except for Massosporda, in which spores
and hyphal bodies are passively dispersed from the disinte-
grating abdomens of the living host cicadas. All of the
diverse types Of secondary spores are also forcibly dis-
charged except for those capillispores (see below), produced
on long, narrow capillary sporophores.

All species of Entomophthora sensu stricto (Remaudiére
and Keller 1980) have campanulate and apiculate primary
spores which are forcibly discharged on a stream of cyto-
plasm and vacuolar sap squirted from the sporophore as the
wall layer(s) securing the primary spore to the sporophore
Break. ine SpOreritsel: Nas a tliat or Sliehtly convex base
rather than a distinct papilla, and appears to play no
active role in the discharge process. Spores discharged in
this manner are accompanied throughout their trajectories

206

by a voluminous drop of the protoplasm which serves to af-
fix the spore to the surface on which it lands, and forms
a characteristic radially striate corona around the pri-
mary spore upon drying.

The primary spores of all species of the non-entomo-
genous genus Bastdtobolus are borne on sporophores with a
pronounced swelling immediately below the primary spore.
Discharge occurs with a circumscissile rupture of the lower
portion of the swelling; the spore and swollen upper por-
tion of the sporophore fly away as the hydrostatic pressure
in the upper portion of the sporophore is released back-
wards like a small rocket (Ingold 1934). Ingold's rocket
analogy is strengthened by the fact that it is often possi-
ble to obtain the firing of a "'second-stage" as -the small
conical projection of the Bastdtobolus sporophore into the
spore itself may evert during flight, thus giving a slight
push against the sporophore fragment and assuring a some-
what longer trajectory for the spore than for the sporo-
phore fragment from which the spore becomes detached.

With the exception of Massospora, all other genera of
the Entomophthorales apparently have primary spores dis-
charged by the eversion of the spore's papilla against the
sporophore (Gallaud 1905; Couch 1939). This mechanism de-
pends upon the establishment of high turgor pressures in
both the sporophore and spore prior to the rounding off of
the turgid cell (Ingold 1971). The sporophore wall breaks
at the point of spore attachment, and the spore papilla
everts rapidly from its original position pointed into the
spore and pushes the spore away from the turgid sporophore.
The shock of this eversion against the pressurized water
column of the sporophore sometimes causes a rupture of the
sporophore tip and exudation of a water droplet at the time
of discharge (@€.g., Page and Humber 1973).

Despite the lack of detailed understandings of these
discharge mechanisms and of the involvements of the various
wall layers of the sporophores and spores, it is apparent
that spore discharge is a major event in the life history
of an entomophthoralean fungus requiring the expression of
a considerable portion of the genome. In view of the sig-
nificance of this process for the fungus and the restricted
distributions of the three diverse mechanisms, it is appro-
priate to recognize the mode of forcible discharge of pri-
mary spores as having taxonomic value at least on the
generic level.

207

MODE OF SECONDARY SPORULATION

The general ability of entomophthoralean fungi to
produce one or more types of secondary spores from primary
spores is one of the most unusual features of this group
of fungi. Among other fungal taxa, the Sporobolomycetac-
eae (heterobasidiomycetous "mirror"' yeasts), many ascomy-
cetes whose ascospores may bud repeatedly in the ascus,
and some other entomopathogens such as a few species of
Septobastdtum Patouillard (Couch 1938) and the hyphomycet-
ous genera Mutogone Thaxter (1920) and Termtartopsts
Blackwell, Samson & Kimbrough (1980) may produce secondary
spores. In none of these groups, however, has secondary
sporulation assumed such a significant role as in the
Entomophthorales. It is becoming increasingly apparent,
for instance, that the secondary spore types are equally
asmantrective -as primary spores or that primary, spores
serve as dispersive units while the secondary spores pro-
duced from them may even be the major infective units
(Carner 1976; Carner and Canerday 1968; Kramer 1980;
Nemoto and Aoki 1975; Nemoto et al. 1979; Selhime and Muma
1966; Tsintsadze and Vartapetov 1976; Wilding 1970).

The morphological diversity (Batko 1974; King and Hum-
Deve ISi)wandebiologicaimimportances of ythese;sporestypes
suggests that they might have taxonomic value above the
Specitrce eve li@um netactietwouty pes. ofesecondaryaspore
forms — microspores and capillispores — have formed the
Dasesmtonitaxonomic divisions at, Subgeneric or generic
levels in the Entomophthorales.

Microspores are produced when the primary spores of
some Contdtobolus and Bastdtobolus species produce a few to
several dozen small secondary spores which are forcibly
discharged and act individually as asexual propagules. The
formation of microspores by C. coronatus (Costantin) Batko
was first used to distinguish the genus Delacrotxta Sacc. §&
Sydow; Tyrrell and MacLeod (1972) proposed that Delacrotxta
should be regarded as a subgenus of Contdtobolus to accom-
modate all species of that genus capable of forming micro-
spores.

Capillispores (despite a profusion of names which are
applied to them) are passively detached spores produced
Singly atop a thin and elongate (capillary) sporophore.
Batko (1966) proposed to divide Zoophthora among four sub-

208

genera wil all species producing almond-shaped to elongate
secondary spores on capillary sporophores to be restricted
to the subgenus Zoophthora Batko. Remaudiére and Hennebert
(1980) restricted the genus Zoophthora to those species
producing capillaries and capillispores and transferred all
species remaining in the other three subgenera proposed by
Batko (1966) to Erynta Nowakowski. In addition to their
presence in all species of Zoophthora subg.Zoophthora sensu
Ben-Ze'ev §& Kenneth (198la) or Zoophthora Batko sensu
Remaudiére & Hennebert (1980), capillispores are produced
by all species of Triplosporium sensu Batko (but not Weozy-
gites sensu Remaudiére §& Keller 1980), Mertstaecrum (inclu-
ding Tabanomyces; see below), and some species of Contdto-
bolus and Bastdtobolus. Batko (1974) considers the pre-
sence of capillary ‘sporophores® to reflect an ancestray
character of the Entomophthorales and rejects the possibi-
lity of thei r=separate orig insmine theses cenerds Dymeovolt-
tionary convergence.

If the Remaudiére classification were prefectly even-
handed: in ats=adoption=of avgenericecri terionspasedson
capillispores, then Entomophthora turbtnata Kenneth (1977),
which produces no capillispores, should not have been in-
cluded in Weozygttes by Remaudiére and Keller (1980), and
both Contdtobolus and Bastdtobolus should have been split..
It is curious that Remaudiére and his colleagues apparently
to not accept the parallel notion of splitting Contdtobolus
because of the presence of microspores in some species of
this genus.

Ben-Zéetev and) Kenneth | (199la)* correctly rejects the
generic emphasis placed on capillispores by Remaudiére and
Hennebert (1980), and propose to use microspores and capil-
lispores as a character with subgeneric validity as did
Batko (1966) and Tyrrell and MacLeod (1972) before them.
However, it is neither advisable nor even possible to apply
criteria based on specialized secondary spore forms to all
genera in which they might occur since Bastdtobolus mtcro-

sporus forms both microspores and capillispores (Benjamin
19625

209

RHI ZOIDS

HUStOn ca lebenspecuLvenands laxonomic, Dist nipuoien

Batko's (1964a-c, 1974) placement of generic value on
the presence of rhizoids has been one of the most trouble-
some aspects of his classification of the entomopathogenic
Entomophthorales. His adoption of this criterion narrowed
his circumscription of Entomophthora s.str. and led him to
disperse the few species now regarded to belong in this
genus (Remaudiére and Keller 1980) among two genera. This
misplaced emphasis also led him to propose Cultetcola and
Entomophaga, separated by the respective presence or ab-
sence of rhizoids, genera whose validity and circumscrip-
Clonsmhavemremainedasuspect, duemtonthe: general controversy
OVermthiesvabidity, of rhazor1dseand) LOmbatkO Seinabii Ety to
provide a definitive separation of Contdtobolus from Ento-
mophthora sensu lato.

Humbersetealmei(1977)mere-evaluated and'rej;ected) the
Significance of rhizoids described for Entomophthora vtru-
lenta Hall §& Dunn (1957; = Conidtobolus thrombotdes Drech-
sler; see Latgé et al. 1980). The rhizoids described for
this fungus are now regarded to have belonged to another
entomophthoralean fungus affecting the same aphids but
which produced no spores or whose spores were undetected.

Remaudiére and Keller (1980) and King and Humber
(1981) independently discuss those species which usually
form rhizoids but occasionally fail to do so. Note that
elmmeunese eXCeNtLONnsarepresents cine: absenccuOb (Ore tarlure
SOutindjmrii zoids in species normally producing «them; they
never involve observations of rhizoids in species not known
to form them. Remaudiére and Keller (1980) believe that no
Weiehtasnouldsbpesputeonsthe presence of rhizo1ds because
PNeitiepTes ence mane cercainespecices. gis NOtwalwaysecOnstanun
On the contrary, King and Humber (1981) regard the presence
of rhizoids to be taxonomically significant while their ab-
sence is not a dependable criterion.* Ben-Ze'ev and Kenneth
(198la) regard the presence of rhizoids to be taxonomically
Significant at the subgeneric level. Brobyn and Wilding
(1977), in the most thorough consideration of the develop-

Brera toi. a lechanvesmotsthestex uniintventronallyaattered
this#opinion tows tate sthateche absence: of rhizoids us
not taxonomically significant.

210

ment of rhizoids and cystidia yet published, uphold the
presence of rhizoids to be a taxonomically significant
character.

It is necessary to consider the general’ situation to
know if any confidence should be placed on the presence
(or absence) of rhizoids as a taxonomic characters. Even
though no absolute scale is available to quantify one's
level of confidence, it is necessary to weigh how well a
species is known: How many times has it been found? In
how many sites? From how many hosts? Are rhizoids always
present or always absent? An examination of the taxonomic
distribution of rhizoids among the entomophthoralean spe-
cies reveals that many have never been found with them
present. In only a few species, particularly Zoophthora
radtcans (= Entomophthora sphaerosperma), is the presence
of rhizoids usual but inconstant. The absence of rhizoids
might be explained in some instances by improper handling
of specimens, but it is also possible that their formation
On some insects might be’ a function of that particular
host (Remaudiére and Keller 1980; King and Humber 1981).

GENERA WITHOUT RHIZOIDS. Rhizoids are completely ab-
sent or have not been found in several genera. “These in-
clude Trtplosportum (at least 7 species), Entomophaga (at -
least 4 species although the circumscriptiom of this genus
remains in dispute), Tabanomyces (1 species; = Mertstacrum),
Massospora (13 species), and Strongwellsea (2 species).

None of the non-entomopathogenic genera — Bastdtobolus,
Aneyltstes, Completorta, Mertstacrum, and Ballocephala —
produce rhizoids.

ENTOMOPHTHORA (SENSU STRICTO). Two of the six species
of Entomophthora s.str. (Remaudiére and Keller 1980) pro-
duce rhizoids. This is the least constant appearance of
this structure in any entomophthoralean genus. It can be
Said with assurance that rhizoids do not occur in the type
species, HE. muscae, since this is the most frequently ob-
served of all species in the order. Both #. cultets (Braun)
Fres. and £. planehontana Cornu produce abundant rhizoids
on hosts bearing forcibly discharged primary spores, but
these anchoring structures may not occur when resting
spores are formed. Both species are cosmopolitan in dis-
tribution and affect a wide variety of dipterans and aphids
(respectively). All of the remaining species — EF. webert
Lakon ex Samson (a little known species from a neuropteran),

E. thrtptdum Samson §& al. (1979; known only from thrips in
Dutch greenhouses), and #. erupta (Dustan) Hall (which is
known from several North American and European sites and
hosts) produce no rhizoids and disperse their primary
spores from living, mobile hosts. The same behavior occurs
in the species of Massospora and Stronguellsea.

CONIDIOBOLUS. Among the 27 morphologically homogene-
OUSmSDCCLCSMLeCOUnIZeadmDY Kinge 10 76a-b,607 /)eenone as
known to produce rhizoids; very few Contdtobolus species
are currently known to be entomopathogenic. Remaudiére and
Keller (1980) transferred most entomophthoroid species with
round to pyriform spores from Entomophaga (which was cir-
cumscribed to exclude species with rhizoids) and those spe-
cies from Cultcicola with round spores to Contdtobolus.

All other Cultctecola species have campanulate spores and
were restored to Entomophthora by Remaudiére and Keller
(1980).

Among the 38 Contdtobolus species listed by Remaudiére
and Keller (1980), four produce rhizoids: Empusa aptculata
ax arena jormanax wake spapt lara. Thax..and LnLomopn—
thora carpentiert Giard (1888). The first three species
were placed in Cultctcola by Batko (1964c-d) because of
enelwerhi Zo dseandmultanucleate round spores borne on ysim-
ple (or occasionally branched) sporophores. Their pro-
nounced similarity to Contdtobolus species belies two obvi-
ous morphological differences from all other Contdtobolus
Species: the consistent formation of a few stout rhizoids
ending in strongly differentiated holdfasts, and the nearly
cylindrical, collar-like narrowing of the sporophore apex.
Close study of these species will be required to verify if
they actually belong in Contdtobolus as suspected by King
(1976b) and affirmed by Remaudiére and Keller (1980).

The only morphological character of Entomophthora car-
penttert Giard (1888) fitting Contdtobolus is the round
primary spore supposed by Turian (1957) to belong to this
Species. Several other characters of this fungus — the
presence ‘of rhizoids,iminucleate spores, on branched sporo=
phores (Turian 1957), and cystidia (Petch 1944, if Petch's
belief that EF. coleopterorwn Petch and #. carpentteri are
identical is correct) — correspond exactly with those of
Erynta or Zoophthora in the Remaudiére classification.
Entomophthora coleopterorum (Petch 1944) has narrowly oval
rather than round primary spores; the mode of rhizoidal at-
tachment is identical with that of F. carpentteri, a spe-

Ale

cies which Giard (1888) characterized only the the distinc-
tive nature of its rhizoidal.attachment. In a situation
resembling that described above for the "rhizoids" of C.
thrombotdes, one cannot discard the possibility that Turian
(1957) may have observed the round spores of a secondarily
invasive Contdtobolus species on a beetle which was already
infected by F. carpenttert which was able to form its very
characteriStic rhizoids but nogspores.. |NOicytolovical
staining or photomicrographic evidence supported Turian's
contention that these round spores were uninucleate; it is
possible that he mistook a,central oil droplet or vacuole
for-arnucleus. The discrepancies between the, Grardseretch,
and Turian concepts of #. carpenttert can be reconciled
only after examining any existing herbarium material of
these collections. In any event, however, the entire habit
of this species refers it to Erynta or Zoophthora rather
than to Contdtobolus.

ZOOPHTHORA.- The presence oF eri zZOLds sUSsest Tone) ear.
related with branched, sporophores bearing yuninuclcate ww. p1—
tunicate primary spores — the species of Zoophthora Batko
or Erynta Nowakowski. All eight Zoophthora species recog-
nized by Remaudiére and Hennebert (1980) form rhizoids. By
including four newly described species, 4. lanceolata Kel-
ler (1980), Z. crassttunteata Keller (1980), Z. ortentalts
Ben-Ze'ev & Kenneth (198la), and Z. petcht1 Ben-Ze'ev &
Kenneth (19381b),. thestotalarisesstorat: leastmulmoteslmspc «
cies known to produce rhizoids. Ben-Ze'ev and Kenneth
(1938ia) remain uncertaim af rhizoids were: presen aonmcue
few specimens of Z. ortentalts available to them.

ERYNIA. Among the species listed by Remaudiére and
Hennebert (1980) and Remaudiére and Keller (1980), some 21
of 25 Erynta species (excluding those of Strongwellsea)
produce rhizoids. Together with Hrynta delphacts (Hori)
Humber*, £. delpintana (Cavara) Humber**, and EF. tthacensts
Kramer (1981), the total incidence of rhizoids rises to 22
of 28 species. The absence of rhizoids in the six Erynta

- ERYNIA DELPHACIS (Hori) Humber, comb. nov., basionym:
Entomophthora delphacis Hori, 1906, Entomol. Mag.
(Tokyo) 8: 81.

** HRYNIA DELPINIANA (Cavara) Humber, comb. nov., basionym:
Entomophthora delpiniana Cavara, 1899, Nuov. Giorn. Bot.
Ita lei Ser aaa? 2

Species discussed below is not considered to overturn the
value of rhizoids as a secondary generic character although
E. delphacts and EF. delptntana constitute the most notable
Ormtiicsceexcep tions.

inexteme( less suspected tnaGrhazoids were ipresent on
Empusa virescens Thax., but could not be certain since he
was not the original collector of the material which he
PA CCTHUSCU mt OPCGSCrIventiismspecies:

Erynta aquattca (Anderson § Anagnostakis) Humber* is
known only from mosquito larvae and pupae floating on the
surface of temporary pools in the northeastern United
states. / This floating habit of the stricken hosts main=
tains their position for favorable transmission of the fun-
gus withouueany » need for rhizo1ds. /alhis species is mot
known well enough, however, to say whether rhizoids might
nOCMrOMN, for cxampley on =individuals trapped atethe
receding edges of rapidly drying pools.

Ben-Ze'ev and Kenneth (1979) report cystidia but no
rhizoids from Zoophthora ertnacea Ben-Ze'ev §& Kenneth, but
note the Similarity of rhizoids and cystidia of species
referable to Zoophthora subg. Erynta Batko (1966). They
muse on the possible inter-relatedness of cystidia and
rhizoids as did Thaxter (1888) and Gustafsson (1965).
However, Brobyn and Wilding (1977) provide effective evi-
dence that the functions and course of development of these
Serle cures sacemtunnelatreduinuspectes wiere sthey mothimoecur
together; it must be assumed, then, that rhizoids and
eystidiasare under completely Separate genetic controls.

Empusa carolintana, with its multinucleate unitunicate
Spores borne on simple sporophores and lack of cystidia or
rhizoids, is once again shown to stand apart from Erynia in

* HERYNIA AQUATICA (Anderson & Ringo ex Anderson & Anagno-
stakis) Humber, comb. nov., basionym: Entomophthora
aquatica Anderson §& Ringo ex Anderson & Anagnostakis,
1980, Mycotaxon 70, 350, NON Entomophthora aquattca
Anderson G Ringo, 1969, J: Invertebr. Pathol%, 73, 386
(which was invalidly published without a designated
type). The combination Hrynta aquatteca (Anderson &
Ringo) Remaudiére & Hennebert, 1980, Mycotaxon 7/7,

30 Pan smal SOmitiva Licdwaccordinget OmiriLcles4>"ousthe
International Code of Botanical Nomenclature.

214

all major characters except for the shape of its primary
spores. The correct generic assignment for this fungus re-
mains problematic, although its spore karyology, sporophore
morphology, and mode of spore discharge more closely re-
semble the species of Entomophaga than any other genus re-
cognized by the Batko, Remaudiére, or Ben-Ze'ev/Kenneth
Clas sutivcat tons.

Erynta delpintana and Erynta delphacts may be true
exceptions to the usually rhizoidal state of fungi in this
genus. Cavara (1899b) is fastidious in his observations of
the small, uninucleate, bitunicate primaryespores., isecon—
dary apores like the primaries, digitately branched sporo-
phores, and very large and occasionally branched cystidia
of FE. delpintana. However, he never specifically mentions
any hyphae affixing the affected flies to the damp piers of
a bridge over a small stream. The flies were completely
covered under a dense, dirty-white to straw-colored hyme-
nium. This entire hymenial covering was easily broken away
from the cadavers during attempts to detach the insects
(Cavara 1899b). Except for the apparent absence of rhizoids
all characters of this species matchjthe scharacteristics
used by describe Zoophthora subg. Erynta Batko (1966); all
species in this subgenus have rhizoids and prominent cysti-
dia. Erynta delptntana is known only from Cavara's origi-
nal collections, but it seems likely that examination of
this material or any fresh collections would reveal the
Presence ofarh@zords:

Erynta delphacts is a much more important exception to
the presence of mhizoi1ds thant thevotherespectesmconsiccred
above. This pathogen of leaf- and planthoppers in Asian
rice paddies bears a strong morphological resemblance to
EZ. neoaphtdis Remaudiére § Hennebert (= Entomophthora apht-
dts Hoffman sensu Thaxter) from aphids (Shimazu 1976, 1977;
Remaudiére and Hennebert 1980), but differs notably in the
complete lack of rhizoids on any of its hosts, including
two artificially infected aphid species, Aphts gossyptt
and Macrostphum akebtae (Shimazu 1977, and personal commu-
Nication). #rynta delphacis produces cystidia both on its
host and in culture. Even though Hori (1906) illustrates
and describes spherical, yellow-white resting spores 24-28
um in diameter and containing many fatty granules, no such
spores have been found in subsequent collections of this
Species in Japan, the Philippines, or Indonesia, nor are
they formed in culture.

pases)

Remaudiére and Hennebert (1980) consider the differ-
ences of growth tm vitro between £. delphacts and E. neo-
aphtdts to be insignificant and regard #. delphacts as a
nomen dubtum. However, a comparison of a Japanese strain
(Shimazu F32; our RS 134) and several Philippine isolates
of E. delphacts (RS 458, RS 459, RS 461, RS 478, RS 479)
from green leafhoppers, Nephotetttx virescens, with numer-
our cultures of £. neoaphtdits isolated by Insect Pathology
Research Unit personnel indicate that #. delphacts grows
substantially more rapidly on a wider variety of media
(nutritionally simple or complex), and sporulates a great
dealmoresprotiitically, overrva Vongerstime sthenvany ‘culture
of E. neoaphtdts we have yet observed. Until more exten-
Sive studies can be made comparing these two species, it
appears to be preferable to recognize H. delphacts as a
legitimate species of Erynia. It should also be noted that
if later studies indicate the synonymy of EF. delphacts and
E. neoaphtdts, the International Code of Botanical Nomen-
clature would require that £. delphacts (Hori) Humber must
be the correct name for the fungus still generally known
as Entomophthora aphtdts Hoffm. (sensu Thaxter).

Much more pertinent information is needed about the
morphology of rhizoids than the perfunctory characteriza-
tion found in many descriptions that rhizoids are merely
“present. \-)sAs noted by “King and Humber (1981) 3 complete
descriptive information about rhizoids should include their
points of emergence, dimensions (particularly thickness),
abundance (few or many?), relative spatial distribution
(isolated, aggregated into one or more pseudorhizomorphs,
or forming a veil?), and the morphology of any terminal
holdfast apparatus. Thaxter (1888), Gustafsson (1965),
Ben-Ze'ev and Uziel (1979), and Ben-Ze'ev and Kenneth
(198la-b) are exemplary in the clarity of their illustra-
tions and characterizations of rhizoidal morphology. This
descriptive information can be useful for detecting dual
infections involving rhizoidal and non-rhizoidal species
Since Brobyn and Wilding (1977) affirm that the morpholo-
Dress OferiilzO1ds) appear to be: speciecsuspecific.

At least three morphological classes of rhizoids can
be recognized: Most rhizoidal species have either (1)
numerous individual rhizoids which are relatively little
differentiated from vegetative and may or may not have any
terminal holdfast differentiation, (2) numerous hypha-like
rhizoids aggregated into one or more columnar pseudorhizo-
morphs, or (3) relatively few, thick rhizoids with strongly

ALO

differentiated (usually discoid) terminal holdfasts. No
comprehensive study of the development and morphology of
rhizoids or of their taxonomic distribution has yet been
undertaken. The thorough studies of Brobyn and Wilding
(1977) were limited to a few species affecting aphids. In
view of the observations presented here, however, such a
comprehensive study would be exceptionally useful.

Rhirzoids as a Taxonomicy Character

The Batko classification adopted the widely accepted
contention of Nowakowski (1883) that the presence of rhi-
zoids is a major generic character. This criterion, super-
imposed on those of nuclear number in primary spores and
the branching of sporophores had effects which have been
regarded as unacceptable and have prevented the general
acceptance of the Batko classification (Remaudiére and
Keller 1980; King and Humber 1981). The Remaudiére classi-
ficatiLonse ine turn. rejected ithetsien7 fi Cancemos at hespre—
sence of rhizoids. It seem certain, however, that the
absence of rhizoids in many genera is significant while
their presence in others is an important secondary charac-
cour

Regardless of the level (if any) at which one may ac-
cept the taxonomic importance -of rhizoidsy species present—
ing exceptions to the normal rhizoidal or nonrhizoidal con-
dition of their genus should be examined carefully. The
classifications of Strongwellsea castrans, S. magna, and E.
earolintana in Erynta and of FE: aptculata, EH. major, E.
paptllata, and FE. carpentterit in Contdiobolus are important
examples of possible misclassifications pointed out by the
lack or possession’ of *rhizoids,

CYSTIDTA

Cystidia (pseudocystidia, paraphyses, or pseudopara-
physes for other authors) are short to long, spear-like to
columnar structures of determinate growth that project
above the level of the sporogenous hymenium. Until recent-
ly, little has been known of their function, but Brobyn and
Wilding (1977) convincingly demonstrated that the cystidia
of Hrynta neoaphtdis are penetration organs which rupture
the host cuticle and aid the subsequent emergence of the

PANG

SPOropnoOres. mene emer gencesOtecystidlda betore the sporo-
phores also has been noted in £. ertnacea (Kenneth 1977)
and £. tthacensts (Kramer 1981). It seems likely that
CVStldlaweservestospertorace tnesnostectticle prior to ny—
menial formation in all species in which they occur; future
investigators should be careful to note the order of emer-
gence of cystidia and sporophores.

Remaudiére and Keller (1980) indicate that, apart from
Erynta and (less commonly) Zoophthora, the only other genus
in which cystidia occur is Contdtobolus. Among the 38 spe-
cies of Contdtobolus accepted by Remaudiére and Keller, the
only ones ever mentioned to form cystidia are C. obscurus,
C. thrombotdes (= EF. vitrulenta), and EF. carpenttert (as E.
coleopterorum Petch 1944); as was noted above, the latter
species almost certainly belongs in Hrynta or Zoophthora
rather than Contdtobolus.

The description of EF. virulenta (= C. thrombotdes;
see Latge et al. 1980) states that “¢ystidia occur rarely”
(Hall and Dunn 1957). However, as explained above for the
mrhiuzords Mor this specresy no ssupsequent treatment Of 1¢
TevULComiiname~cl (nereriuzOLds@eOr CYStidias mine CyYSstt.
dia" of C. thrombotdes might have been those of another
entomophthoralean fungus which produced both rhizoids and
cystidia (see Humber et al. 1977). Hall and Dunn (1957)
might also have interpreted vegetative or sporogenous
hyphae whose indeterminate growth occasionally far exceeds
enewveneraleicevel, Ore tie nymen1im LONbe cySticia,.

Remaudiére et al. (1979) report the formation of long
cystidia by C. obscurus on aphid cadavers placed in a humid
environment. I have observed similar growth of long hyphae
above the hymenium of aphids affected by C. thrombotdes but
always find these structures to be undifferentiated vegeta-
tive hyphae which should not be regarded=as cystidia by the
definition given above.

The listing of Lamta (= Entomophthora) cultcts in a
chart by Nowakowski (1883) as having cystidia present and
rhizoids absent was corrected by Batko (1964c) as being a
typographical transposition not corresponding with the Pol-
Ponerextawit CirmcOLrectly mnOteSmcii Ss sSpeClesico NaVvemrnii—
Zo1ds DUE NO cystidia.

The genus most closely related to Zoophthora and
Erynta must be Stronguellsea, with its uninucleate and bi-

218

tunicate primary spores borne on simple rather than
branched sporophores (Batko and Weiser 1965; Humber 1976).
Remaudiére and Keller (1980) consider the lack of rhizoids
and cystidia to be insignificant criteria and use this in-
formation to explain their submergence of Stronguellsea
into Erynta (see Humber 198la). In fact, if cystidia func-
tion only for cuticular penetration, it is not surprising
that Strongwellsea species form no cystidia. These fungi
produce no hymenial structures outside the host's body.
The abdominal hole (the only cuticular rupture caused by
the fungus) begins with a point perforation of the pleuron
and enlarges progressively. This point initiation of the
rupture appears to be involved with the hypertrophic reac-
tion of the host cuticle which leads, in turn, to the ex-
tensive lining of the lower portion of the fungal ball by
the host's cuticle. It seems clear that some mechanism
completely distinct from that of cystidial penetration is
involved in the formation by Strongwellsea of an abdominal
hole (see Humber 1975, 1976).

In view of the above discussion, it srs probably jcor-
rect to say that the only species in which cystidia (as
determinate organs for cuticular penetration) occur are
those with uninucleate, bitunicate primary spores borne on
branched sporophores and classifiable in Erynta or (less
commonly) Zoophthora sensu Remaudiére §& Hennebert.

Unlikesforerhazoids,. thespresencesorgabcencomOtecy oti.
dia has never been used to separate genera although the
Batko classificaiton recognized their value as a supporting
character strongly correlated with the occurence of uninuc-
leate, bitunicate primary spores and branched sporophores.
ASawith the other charactersediseussedshere, sexceptlousaco
the usual state of thescharactersinga cenuseshouldgindicave
thateae close sinspecctonsormt ne cOLTespOnucncesmO Leda lst
characters of the suspect species with those of other spe-
ciespin they genusuiseinvorder wealnesp resencemOcacystldl amin
E. coleopterorum (Petch 1944), a presumptive synonym of
Contdtobolus carpenttert (Giard) Remaudiére § Keller, of-
fers a significant example of this principle.

219

RESTING SPORES

aygospores or Azygospores:
A Morphological/Cytological Dilemma

I applaud the de-emphasis by Remaudiére and Keller
(1980) of the distinction between zygospores and azygo-
Spores. sineyenotercorrectlys that resting Spores are not
known (or are not formed) in a series of species, and that
the mode of their formation has not been observed in a
Parsee propored Onsotaspecics sctia badQsproducemsres ti ngEspores,
Both types of development may occur in closely related spe-
cies or even within a single species (MacLeod and MUuller-
Kogler 1973; Humber 1976; Remaudiere and Keller 1980; King
and Humber 1981).

This developmental distinction has been used only
twice as a taxonomic character of generic importance. The
peculiar mode of conjugation of (amoeboid?) hyphal bodies
or gametangia of Empusa fresentt Nowakowski 1883 (= Neozy-
gttes aphtdis Witlaczil 1885) and the resultant jet-black,
ovoid zygospores formed at the point of conjugation prompt-
ed Ihaxter (1888) to set this and related:species apart iin
Empusa subg. Tritplosportum. This taxon was later raised to
generic status by Batko (1964b), who was apparently unaware
of the nomenclatural priority of the generic name Neozy-
gttes Witlaczil (see Humber et al. 1981). King (1976b)
notes that the double-walled resting spores formed by the
species of Contdtobolus are anisogamous zygospores. Other
types of conidioboloid resting spores such as chlamydo-
spores and villose spores are distinguished from zygospores
(or azygospores) by their single rather than double wall
SCRUCTULeS.

Couch (1939) notes that the zygospores of Contdtobolus
species are formed in the larger of two gametangia (and are
almost always produced in the axis of the parental hypha)
while those of Entomophthora (sensu lato) are produced in
lateral buds from the conjugating hyphae or hyphal bodies.
Very few species of Entomophthora s.1. produce their zygo-
spores in the hyphal axis: FE. vtrulenta (= C. thrombotdes),
E. obscura Hall & Dunn [= £. tgnobtlts Hall §& Dunn = #£.
thaxtertana (Petch) Hall §& Bell; see below], and Empusa
diptertgena Thaxter. The former two species are now recog-
nized as species of Contdtobolus by Remaudiére and Keller
(1980). With the apparent exception of £. dipterigena,

220

whose zygosporogenesis is illustrated by Gustafsson (1965),
the resting spores of other species of Entomophthora or its
segregated are budded off from the parental hypha or hyphal
body.

Zygospores, the bilayered, thick-walled spores derived
from a conjugation of two gametangia, have been a well
established phenomenon in the Entomophthorales since their
first recognition (Brefeld 1973; Nowakowski 1877, 1882,
1883; Thaxter (1888). Nearly every subsequent major discus-
sion of these fungi has referred to zygospores as sexual
spores and to azygospores as asexual spores (Schroter 1897;
MacLeod 1956, 1963; Gustafsson 1965; Waterhouse 1973).

In general morphological terms, zygosporogenesis may
be called a sexual process, in which case azygosporogene-
Sis must be regarded as asexual. However, in their strict
sense, the terms zygospore and azygospore refer only to the
mode of development of a thick-walled spore in the Zygomy-
cotima (Fitzpatrick 1950; Bessey #1950 Ainsworth ol:
Snell vand Dicks 19713" AlexopoulosmandsMims se! 979)e.

There is a hazard in’ thinking sotm@zycosporeseas sexual
and azygospores as asexual: One can be misled easily into
performing an act of semantic sleight-of-hand by then ex-.
pecting that the genette events of a sexual life cycle,
karyogamy and meiosis, necessarily occur in a ''sexual"
zygospores but cannot occur’ in an “asexual” azygosporey A
related terminological confusion can occur when one refers
to the resting spore as the sexual spore in the entomoph-
thoralean life “cyclesasvopposedmto sthestorcib lyedrscharced
primary -(or*secondary) @spores which ane, anseveryeasense-
asexual spores.

Nuclear events in entomophthoralean resting spores —
reductions in nuclear number, karyogamy, and meiosis —
remain completely uninvestigated in all but a few species.
This gap in our knowledge reflects a lack of fundamental
Studies on the life cycles’ in’ this group.

Some cytological studies have noted the progressive
reductions of nuclear numbers to two or even a single
nucleus in fully mature resting spores (Vuillemin 1900a-b;
Krenner 1961; Latgé 1976; Couch et al. 1979; D. McCabe and
B. Martinell, unpublished). Other investigators using
different species have found no obvious change in the
nuclear number from the time of formation to the fully

TOE

mature, dormant state (Olive 1906; Riddle 1906; Goldstein
1929; Humber 1975). If the Entomophthorales follow pat-
terns of karyogamy and meiosis similar to those in other
Zygomycetes (Cutter 1942a-b), karyogamy should occur in the
resting spore followed by meiosis in the spore just before
or during its germination. No conclusive cytological evi-
dence of nuclear fusions or of meiotic division has yet
been found in entomophthoralean fungi despite the strong
circumstantial evidence for’ their occurences (e.g¢., Couch
tera dee 97.98): It must be hoped that synaptonemal com-
plexes, a widely recognized ultrastructural marker for
chromosomal synapsis, will be demonstrated at the time and
ST LCeOMeticapresumpulVve mclOsi se inmsoncsOrsmoresmelners, of
the Entomophthorales.

Data regarding changes in nuclear numbers presented
by Latgé (1976) support the possibility that karyogamy and
meiosis occur in the resting spores of C. thrombotdes, and
tiatecnesvevetatiVesnuclei, OL thisespecies must pe haploid,
Albeit, Latgé feels that his data are also compatible with
gametangial meioses of diploid nuclei such as occurs in the
Oomycetes (see Alexopoulos and Mims 1979). Little ambigu-
Peyeaboucethc interpretation OL tice! lilcecyCcie.remains,
however, in view of the results of electrophoretic analyses
of isozyme patterns of several strains of C. thrombotdes
and Z. raditcans (May et al. 1979) and of EF. muscae, Masso-
spora etcadina Peck, and Entomophaga gryllt (Fres.) Batko
(Soper, May,@and*@Martanell; invpreparation): These results
Suggest that the vegetative nuclei of these species are
haploid, although heterokaryosis may occur in some species
Si ncemmuluipleralleles;otrsome loci may, bespresent. inva
mycelium.

There is no evidence yet of heterothallism within the
Entomophthorales. All zygosporogenesis appears to be homo-
thalliceandmleavess opens atileaststwo possiblesinterpreta=
tions torstheo life cycles of these: fungi: Those species
whose zygospores or azygospores at some point contain only
a single nucleus probably have autogamous (sexual) life
cycles. Those species whose zygospores or azygospores
undergo no reduction in nuclear numbers may have abandoned
a sexual life cycle in favor of an apogamous (asexual) life
which preserves the particular (successful) combination of
characters in that species. No evidence now exists that
parasexuality occurs in any species of the Entomophthor-
ales.

Zee

The distinction between zygospores and azygospores
cannot be retained as a character of taxonomic signifi-
cance although the presence and nature of zygospores is a
useful ancillary character in at least Triplosportum and
Contdtobolus. The developmental origins of resting spores
can be difficult to interpret correctly even itesuitapie
material is available. Nuclear events in resting spores
are of far greater importance for the fungus than whether
conjugations precede sporogenesis. Karyogamy and meiosis,
the critical genetic events in a sexual life cycle, appar-
ently may occur in either zygospores or azygospores;
similarly, other species may have apogamous life cycles
regardless of whether their resting spores are zygospores
or azygospores.

Indeed, the full details of entomophthoralean life
cycles and their breeding strategies may yet prove to be
useful taxonomic characters, but not enough is currently
known of these processes to draw meaningful conclusions.
There is no doubt, however, that this information is of
absolutely vital importance for the practical development
and manipulation of these fungi for use in the microbial
CONtTLO MMOL Sethe reinsectanostse

Resting Spores: Problems and Interpretation

of Taxonomic Traps

It appears that the generic events of sexual repro-
duction might occur in either zygo- or azygospores, and
that even species producing ''sexual"' zygospores might be
apogamous. Several further caveats and an example serve to
reinforce the rejection of any taxonomic value which might
be attached to the distinction between zygospores and
azygospores (Remaudiére and Keller 1980).

In most instances, authors simply state their obser-
vation that the resting spores of a given species are
zygospores or azygospores and neither illustrate nor dis-
cuss the evidence supporting this conclusion. However,
Sawyer (1931) notes the great difficulty in determining if
conjugations occur in Z. radtcans. Humber (1975) also
notes difficulties with the resting spores of Stronguellsea
Species due to problems of interpreting the morphology of
some hyphal bodies and to the evanescence of the emptied
hyphal wall remnants which provide the only temporary evi-
dence for the mode of formation of mature resting spores;

determining the 'pedigree™ of these resting spores is
dependent upon seeing them during a relatively narrow
Periodsuuting their development. Consequently, 1t seems
unwise to place full confidence in any characterization of
resting spores as zygo- or azygospores unless convincing
documentation is also presented.

The difficulty in correctly applying these developmen-
tal designations casts doubt on whether any taxonomic de-
cisions should be based solely on this distinction (even
though it may haye some utility for characterizing, but not
Separating, some genera as noted above). Ihe practical ef-
ECCUS Ompditrering ViewpOINtseorstice ValuceOLt thts distinc-
tion can be seen in the following example:

In studies on the identities of aphid-pathogenic
Species of the Entomophthorales, Humber (1978) noted that
Entomophthora tgnobtits Hall & Dunn should be used as the
nomenclaturally valid name for Entomophthora thaxtertana
(eeccijelial wambelipeaespecleselouwhich Hal leand Bell
(1963) considered F. tgnobtlts to be identical. Empusa
thaxtertana Petch (1938) was proposed for a species mis-
identified by Thaxter (1888), and which was characterized
as having azygospores. The resting spores of E£. tgnobtits
are described by Hall and Dunn (1957) as zygospores. Hum-
ber (1978) considered the reported differences in the ori-
gins of resting spores to be so ambiguous as to be insigni-
ficant, an opinion which is reinforced here. A later-study
by Remaudiére et al. (1979) used morphological and bio-
chemical data to demonstrate the synonymy of EF. thaxtert-
ana and EF. obscura Hall §& Dunn (1957) [= Contdtobolus ob-
seurus (Hall & Dunn) Rem. & Keller], which was described
as producing azygospores.

In fact, the differences between EF. thaxtertana, E.
tgnobtlts, and E. obscura had always been troublesome for
diagnosticians. The only major described difference be-
tween #. tgnobtlts and HF. obscura is that of zygospores
versus azygospores, a difference considered here, by Humber
(1978), and by Remaudiére and Keller (1980) to be insigni-
ficant. A more consistent and simple approach to this
taxonomic problem would have been to accept EF. obscura to
Hestieecorrectundice DyaVvilTtucrOL pacenpriOrityeoveras.
tgnobilts rather than to regard the latter species as a
nomen dubtum merely because of the described difference in
the, ortegins of the resting spores in these two species
(Remaudiére et al. 1979).

224

If there is to be even-handed application of the
accepted taxonomic criteria and acknowledgement of the net
effects of rejecting other criteria which have been applied
in the past, it iS now necessary to recognize Unescorrect
synonymy of Contdtobolus obscurus to be the following:

Conidiobolus obseurus (Hall & Dunn) Remaudiére §
Keller 1980, Mycotaxon 17, 331.

= Entomophthora obscura Hall.& Dunn 1957, Hil-
GAraGHae: 7,902.

= Entomophaga obscura (Hall & Dunn) Batko 1964,
Bulle gAcad. Polons Cia ocr OCummD Lois
12, 404.

Entomophthora tgnobtlts Hall & Dunn 1957, Hil-
SardiauaieeloZ.

Entomophthora planchontana Cornu sensu Thaxter
1888, Mem. Boston Soc. Nat. Hist. 4, 165,
NON Sensu. Petch 1948 y9hrans bri teeny COlemoOG.
Pil a,

= Empusa thaxtertana Petch 1938, Trans. Brit.
Mycol. Soc. 21, 34. NAME INVALID WITHOUT
LATIN DIAGNOSIS.

[= Entomophthora thaxtertana (Petch) Hall §
Bell 196355 “Jesinsect Pathol. worm Usoe|

[= Entomophaga thaxtertana (Petch) Batko 1964
Bulle Acade RPalon!aeSer soci aabLOl amie
404. ]

CHOOSING A CORRECT CLASSIFICATION:
TRIAL BY ANOMALY

One of the most severe tests for a classification
SystemersetOusce NOWaner tm GecalanalGl oeticulcast suyl teal
species from the group being classified. Withanethe, Ento-
mophthorales, several seemingly anomalous generic assign-
ments from both the Batko and Remaudiére classifications
are discussed here. Howeverj;iin terms of testing the real
capacities of these or any other classifications of these
fungi, confirming the correlations of characters, or sug-
gesting where primary taxonomic weights should be placed,

Dab

the most important fungus has not yet been considered.
PitSetseareunuescriped=species tound Dy sbadazy 31975) ein
Poland which was, obviously, not known at the time of the
proposition of the Batko classification, and was not con-
sidered by the Remaudiére classification.

THiswrungusmathixcceolatsmiliptera.=lendipeataae) aby
rhizoids to wet, decayed wood just above a stream surface
in a deciduous forest in the Wielkopolski National Park,
Poland. Its primary spores are Slender and curved, 45-75
(82) x 8-10.5 um, strongly tapered apically and with a
frat cencdmm conical spapi | laemeeinesesciaraclers: a1mostsexace-
ly match those of Erynta contca (Thax.) Rem. §& Henn., which
RaSmcucCndetEClLVCdusPOLCSmlOLedmsUZCE LOCH El Ca ml eDatazy) Ss
fungus) with bitunicate walls and single nuclei, borne on
branched sporophores, and with cystidia and rhizoids pre-
Sent. .batazy's fungus) 1s, however, clearly different from
E. contea since its primary spores have unitunicate walls,
contain 4-10 nuclei, and are borne on simple sporophores ;
cystidia are absent.

The Remaudiére classification, with its primary empha-
sis on spore morphology, would place this species in Erynta
even though that assignment would be impossible if one ac-
cepts the emended circumscription advocated here (excluding
Stronquellsea and EL. caroltntana from the genus by limiting
ite COUSPEC1ES With uninucleate, bitunicate spores: borne .on
branched sporophores and not producing capillispores). The
Batazy fungus might be regarded as a species of Cultctcola
except that Batko (1974) apparently rejected Cultetcola as
a heterogeneous mixture of species now regarded to belong
to Entomophthora s.str. or Contdiobolus (Remaudiére and
Keller 1980; King and Humber 1981). There is no basis for
including the Batazy fungus in Contdtobolus according to
Phescircumscripiionssot ethicmeenusecithenestated ors implied
by Batko (1974), Remaudiére and Keller (1980), here, or in
Humber (1981b). It might be assumed that the Polish fungus
could be allied to EF. aptculata, EF. major, and E. paptllata
becauscuoL their multinucleate uni tunicatesspores wexcepe
Pnatetneanuclearsmorphotogysotethe slat tersspeciesscorre-
sponds to that of Contdtobolus species while that of the
Polish fungus resembles Entomophthora and all of its segre-
gates rather than Contdtobolus (see Humber 1981b). The
generic system outlined by Ben-Ze'ev and Kenneth (1981la)
would include this species in Entomophthora sensu stricto,
but the mode of spore discharge in the Batazy fungus is by
Ciceeversi Oneotethe papilla:

226

If, as suggested here, one rejects spore morphology as
thei primarysgeneric criterion and accepts the restriction
of Erynta noted above, and places generic value upon the
karyology of primary spores, the morphology of the sporo-
phore, and the mode of spore discharge, the only remaining
sensible disposition for Batazy's fungus is in a new and
(for now) monotypic genus allied to but differing from
Erynta. It seems reasonable to assume that other entomoph-
thoralean species may yet be found with characters like
those of the one characterized by Batazy (1978).

It is imperative that this Polish fungus be fully
characterized and more widely publicized. This fungus
seems certain to provoke much useful discussion of the
taxonomic criteria which should be accepted for entomoph-
thoralean fungi, and of how much weight should be accorded
ComeacheOLathescacricenlar

MODERN VIEWS OF ENTOMOPATHOGENIC
GENERA OF THE ENTOMOPHTHORALES

This study has examined the major morphological cri-
teria used in entomophthoralean taxonomy, particularly as
they are applied in the generic classification schemes pro-
posed by Batko and by Remaudiére and his colleagues. It is
apparent from all of the above considerations that the
application of only three characters provides an effective
separation of species into morphologically and biologically
homogeneous genera. These characters are the following:

1. The number and nature of nuclei in
the primary spores.

Zo Ines morpho logysobathessporophores
(and the mode of sporogenesis).

3. The mode of discharge of primary
spores.

Neither the morphology of primary spores, types of
secondary spore formed, presence or absence of rhizoids,
presence or absence of cystidia, nor the mode of formation
of resting spores was found here to be important for de-
limiting genera in the Entomophthorales. Each of these

Vogt

characters, however, may have utility as secondary or sup-
porting characters which do not in themselves determine the
PimVceSwOL sscicra = so oOme OL these Characters arevappropriate
for the circumscription of subgeneric taxa (Ben-Ze'ev and
Kenneth 198la).

It is appropriate to summarize the difference among
the Batko and Remaudiére classifications and that proposed
preliminarily by Ben-Ze'ev and Kenneth (198la), and to note
those adjustments which seem to be necessary to circun-
Scribe these genera still more accurately according to the
Gr iecTiasaccep ted ene:

ENTOMOPHTHORA Fresenius. Batko's overestimation of
the importance of rhizoids caused him to split a small and
natural group of species (MacLeod et al. 1976; Samson et
al. 1979) with simple sporophores, campanulate apiculate
primary spores containing a relatively small number of nu-
clei, and a characteristic mechanism of spore discharge
among two genera. The application of the mechanism of
spore discharge as a generic criterion confirms the narrow
definition of Entomophthora proposed by Remaudiére and Kel-
ler (1980) despite the de-emphasis here of the value of
spore morphology. The exceptionally high degree of morpho-
logical and biological homogeneity of this group of species
would be manifestly lowered if the mode of spore discharge
did not exclude both Empusa caroltntana Thaxter and the
species found by Batazy (1978) and discussed above. Both
of these species have simple sporophores and plurinucleate
primary spores but are discharged by papillar eversion
rather than by the "fungal cannon" mechanism of #. muscae.

MASSOSPORA Peck. This venerable genus which affects
gregarious cicadas (Homoptera: Cicadidae) was not treated
in the Batko classification until his phylogenetic treat-
ment (Batko 1974) since Massospora represents the only
genus of the entomogenous Entomophthorales whose validity
and circumscription (see Soper 1974, 1981) has never been
questioned. The species of this genus have plurinucleate
primary spores which are not forcibly discharged from the
Simple sporophores lining small cavities in the abdomen of
affected insects.

TRIPLOSPORIUM (Thaxter) Batko, nom. gen. conserv.
prop. sBatkow(1964b) taccepted Whaxter"s (18838)) belief ithat
E. fresentt and similar species should be accorded generic
Status. This small and natural group of species is. distin-

228

guished primarily by its unique mode of zygosporogenesis
and ovoid zygospores with black (or very dark) epispores.
Batko (1964b) circumscribed the genus to include species
whose primary spores are quadrinucleate, borne on simple
sporophores, and capable of producing capillispores as one
type of secondary spore. Remaudiére and Keller (1980) ac-
cept Batko's circumscription of this genus but they (1)
reject the requirement for the primary spores to be quadri-
nucleate in order to justify the (contentious) inclusion of
E. turbtnata Kenneth (1977), and (2) replace Trtplosportum
(Thaxter) Batko with the older generic name Weozygttes Wit-
laezil (1885) Humber ct als (i98l))" otter severaiereasons
why Trtplosportum should be conserved against the older and
nomenclaturally correct name Neozygites; this nomenclatural
issue will be resolved at the 13th International Botanical
Congress during the summer of 1981.

ENTOMOPHAGA Batko. This genus, based on Entomophthora
grylit Fres., was proposed to include species wtih multi-
nucleate spores and simple sporophores but without forming
rhizoids. Its validity has been questioned because of its
linkage to the unacceptable Cultetcola through Batko's
emphasis on rhizoids and also because no satisfactory cri-
terion has been proposed to distinguish unambiguously be-
tween any of these species and the morpholgoically similar
Contdtobolus. The lack of this criterion seemingly prompt-
ed Remaudiére and Keller (1980) to emphasize spore morpho-
logy by transferring all species of Entomophaga and those
of Cultetcola with round rather than campanulate priamry
spores to Contdtobolus.. However, the criterion of nuclear
morphology noted above and discussed fully by Humber (1981b)
readily delimits Contdtobolus from Entomophthora sensu
Stricto and its segregate genera, and confirms the validity
of Entomophaga for species with multinucleate unitunicate
primary spores containing large, readily stainable nuclei,
and discharged by papillar eversion from simple sporophores.

ZOOPHTHORA Batko and ERYNIA Nowakowski. Batko (1964b)
described Zoophthora, with Entomophthora radtcans Brefeld
(= Entomophthora sphaerosperma Fresenius) as its type, for
all species having uninucleate, bitunicate spores borne on
branched sporophores and producing rhizoids. This genus
has been accepted as a natural and valid grouping. Four
subgenera were proposed by Batko (1966) based on the
morphology of sporophores, rhizoids, and cystidia, and the
type of secondary spores produced. Remaudiére and Henne-
bert (1980) limited Zoophthora to species capable of form-

229

ing anadhesive secondary spores atop capillary sporophores
(see King and Humber 1981), with all other Zoophthora spe-
cies being transferred to Erynta Nowakowski (1881). The
Remaudiére classification modified the sense of Batko's
grouping to accommodate species with simple sporophores and
even multinucleate unitunicate primary spores so long as
the spore shape was similar to that in other Erynta
SDSCLES

For purposes of discussing the Remaudiére classifica-
Cionmsanesthisepaper, at has been necessary to accept this
generic separation on at least a provisional basis. Con-
siderable objection can be raised, as noted above, about
ehesuserotecaprl larvesporophores ease ar charactemsom generic
Send ol Cancen mais sSuemoasmDecns treated pant by ,Ben~
Ze'ev and Kenneth (198la) and will be discussed more fully
Dymtnese@autnlors@ineassubsequcnts paper.

A nomenclatural problem reminiscent of the simultane-
ous use of Empusa Cohn and Entomophthora Fresenius as taxo-
nomically distinct genera has been raised by Remaudiére and
Hennebert's use of both Zoophthora Batko and Erynta Nowak-
owski. In 1881, Nowakowski proposed the genus Lrynta to
include Entomophthora ovtspora Nowakowski (1877) and Ento-
mophthora curvispora Nowakowski (1877), but later rejected
this generic name in favor of Brefeld's (1877) usage of
Entomophthora as taxonomically distinct from Zmpusa (Nowak-
owski 1882, 1883). At the time of its description, Zooph-
thora Batko (1964b) included only its type species, Ento-
mophthora raditcans Brefeld. However, the inclusion in
Zoophthora of FE. ovtspora and FE. curvtspora (Batko 1964d)
automatically required the adoption of the earlier name.
Batko (1966) was incorrect in believing that Nowakowski's
later disuse of Erynta removed that name from consideration
in matters of nomenclatural priority.

The ultimate resolution of this nomenclatural problem
with Erynta and Zoophthora depends on the outcome of the
debate to establish a concensus opinion of whether the Bat-
_ko or Remaudiére classification, or some modification of
one of them shall be accepted for the species with bituni-
cate, uninucleate spores on branched sporophores. If, on
the one hand, two genera are recognized with one based on
Entomophthora ovtspora, the type species chosen for Erynta
by Remaudiére and Hennebert (1980), and the other on Ento-
mophthora radtcans, there is no nomenclatural problem to
resolve. If, on the other hand, common practice rejects

230

any generic separation and recognizes only a single genus
which includes both £. ovtspora and E. radtcans, then
either Hrynta would have to be adopted as the correct gene-
ric name or Zoophthora might be proposed for conservation
against Hrynta.

It is unfortunate that such a confusing nomenclatural
issue should arise once more in the Entomophthorales, but
the three possible resolutions are unambiguous. My person-
alsopinaon should tbe 'cleareirom this spapey. me lecOsilouEpe-
lieve that the species originally classified by Batko in
Zoophthora should be split between two genera separated by
characters such as spore morphology and the formation of
capillary sporophores. As much as it might prevent some
amount of further confusion in the literature if Zoophthora
Batko were conserved against Hrynta Nowakowski, I do not
believe that such a proposal would be accepted for incorpo-
ration in the lists of nomina conservanda in the Interna-
tional Code of Botanical Nomenclature. The great prepon-
derance of species proposed as or transferred to Zoophthora
have already been transferred to Erynta by Remaudiére and
Hennebert (1980) or here; the relative number of species
required to be transferred as a consequence of conservation
of a younger name against an older and nomenclaturally cor-
rect one appears to be one of the major concerns of the
nomenclatural committees which decide these matters. The
new combinations in Hrynta proposed here are fully justi-
fied and nomenclaturally correct whether one accepts the
taxonomy of Batko (and his followers) or or Remaudiére and
Hennebert for classityinosthe species sinsdquesuion=

STRONGWELLSEA Batko § Weiser emend. Humber. Humber
(1976) provided an emended generic description and valida-
tion of this genus whose spores are uninucleate and bituni-
cate but are borne on simple rather than branched sporo-
phores, and are discharged by papillar eversion. Remau-
diére and Keller (1980) provided no effective rationale
for rejecting the importance of sporophore morphology or
other diverse supporting data (Humber 1976) in order to
synonymize Strongwellsea with Erynta. There can be little
doubt from the criteria considered here to be significant
at the generic level that Strongqwuellsea must be recognized
aS vamseparate genus.) -Asmore extensive; support stores enls
Opinion is presented by Humber (198la).

Zor

CULICICOLA Nieuwland. This genus is nomenclaturally
superfluous since its type species, Entomophthora cultects
(Braun) Fres., belongs in Entomophthora sensu stricto
(Remaudiére and Keller 1980; King and Humber 1981). The
species included in this genus by Batko (1964c-d) are now
dispersed to Entomophthora s.str. or Contdtobolus Brefeld
by Remaudiére and Keller, although some question remains
whether the conidioboloid species producing rhizoids —

E. aptculata, E. major, and FE. paptilata (which was not
classified by Batko, 1964d, but clearly belonged in this
group) — should be in Contdtobolus or in a closely allied
but different genus. Batko's inclusion in Cultctcola of
E. vtrulenta (= C. thrombotdes; see Latgé et al. 1980) was
based on the erroneous description of this species as pro-
ducing rhizoids (Humber et al. 1977).

Batko regarded this as the most tentative of his pro-
posed segregate genera, and he later apparently rejected
Culteteola in his extended justification of his taxonomic
ideas and discussion of phylogeny in the Entomophthorales
(Batko 1974). Cultetcola has been the most objectionable
of hisegenera tor other studients, of these«fungi1.

CONIDIOBOLUS Brefeld. Even though Batko and Weiser
(1965) note that resting spores of Contdtobolus are not
budded off to the side as in all other genera considered in
the Batko classification, they again cited the widely used
but untenable "'criterion'' which supposes Contdtobolus spe-
SLeSetoubeabasicallyesaprobesawhide, those) of the other
genera treated were entomopathogenic. Remaudiére and Kel-
ler (1980) regard the lack of a suitable criterion to de-
limit Contdtobolus from morphologically similar species put
in Cultetcola and Entomophaga by Batko (1964b-d), with
round to pyriform, multinucleate primary spores and simple
spores, to justify their inclusion in a broadly redefined
Contdtobolus.

As indicated here, however, nuclear cytology does pro-
vide the criterion sought to delimit Contdtobolus species
from those of Entomophthora and its segregates (Humber
1981b). Some of the species transferred to Contdtobolus
from Entomophthora sensu lato truly are species of Contdto-
bolus; these include £. virulenta (= C. thrombotdes; Latgé
et al. 1980), C. obscurus, and possibly C. aptculatus, C.
majon,,and.C. papiilatus (although these last three’ species
may belong in a separate genus allied to Conidtobolus due
to marked differences in the structure of the sporophore

Boz

with the presence of rhizoids as a supporting character).
Other species — e@.g., EF. gryllt, FE. batkot, and EF. gigan-
tea Keller — have nuclei whose morphology indicates their
closer affinities to Entomophthora than to Contdtobolus
(Humber 1981b).

TABANOMYCES Couch, Andreeva, Laird, & Nolan. This
little known genus (Couch et al. 1979) was described for
the peculiar mode of germination and spore formation from
entomophthoralean resting spores found in larvae of tabanid
flies in the Soviet Union.; The thick-walled; coloriess,
ovoid resting spores of the type species, T. milkot (Dudka
& Koval) Couch §& al., were originally described as Coelomo-
myces mtlkot Dudka & Koval. During the germination of the
spores, the single nucleus undergoes two’ (meiotic?) “divi-
Sions, and a short, thick, quadrinucleate sporophore forms.
This hyphasbecomes septate eandecachwuninuctecates cel o-
duces and discharges a lateral primary spore which is, in
turn, capable of producing a secondary spore on a short
Capillary =sporophore (Couchmet 417] 970) neseuevenecmalic:
structures are essentially identical to those described for
the germination of resting spores in the nematophagous
genus Mertstacrum Drechsler (Davidson and Barron 1973).
ThesnucleiallustratedebpyeCouchectealem Gloy ljmarce col ative>
ly small and have a prominent central nucleolus; this type
of nucleus resembles that of Contdtobolus (Humber 1981b)
and Mertstacrum (Humber, unpublished).

Remaudiére and Keller (1980) do not mention Tabano-
myces, but this genus cannot be easily placed in any entomo-
genous genus included in the Batko or Remaudiére classifi-
cations. The remarkable similarity of Tabanomyces and
Mertstacrum indicates that these genera are synonymous and
that 7. milkot must be recognized as a heretofore unknown
entomopathogenic species of Mertstacrum*.

TARICHIUM Cohn. This genus was created for those spe-
cies of the Entomophthorales known only by their resting
spores (see MacLeod and Muller-Kogler 1970). Neither the

* MERISTACRUM MILKOI (Dudka §& Koval) Humber, comb. nov.,
basionym: Coelomomyces mtlkot Dudka §& Koval in Dudka,
Koval § Andreeva, 1973, Novitates Systematicae Plant-
arum non Vasculartum 10, 88-91. SYNONYM: Tabanomyces
mtlkot (Dudka § Koval) Couch, Andreeva, Laird § Nolan,
197975 Proce Natle Acadn ocr (USA) «cre 2090-500

255

Batko nor Remaudiére classifications fully accept this
genus, but properly regard its members as having still un-
determined affinities. It can be assumed that Tartchtum
Species may be connected to a species producing primary
spores and, thus, placed in the appropriate modern genera.
A taxonomic system which accepts a narrow sense of knto-
mophthora and a number of other entomopathogenic segregate
genera must also accept Tartchtum as the correct provision-
aieCeneric nane stOrethe specres involved ratner than the
name Entomophthora (Tartchtum) as was suggested (without
any formal nomenclatural status as a subgenus of Entomoph-
thora) by MacLeod and Muller-Kogler (1970).

Concluding Remarks on Generic Classifications

Despite its manifest shortcomings, the classification
proposed by Batko (1964a-e, 1966; Batko and Weiser 1965)
forms the basis for all other contemporary approaches to
a generic classification for the Entomophthorales. This
classification is discussed in depth (although not substan-
Cia tbyeanp iit ed )ymeitimecicurirstatcempt CO OUGLine) Ee
course of evolution in the Entomophthorales (Batko 1974).
The Remaudiére classification attempted to eliminate the
flaws which Remaudiére and his colleagues perceived to pre-
VentecncmcciCTraleaccepuancesO1sthespackO Classit) Catton amen
generic arrangement more similar to that proposed by Batko
than by Remaudiére, Hennebert, and Keller emerges here in
view of the acceptance of primary spore karyology, sporo-
phore morphology, and the mode of primary spore discharge
as the three characters best suited for delimiting genera.
Ben-Ze'ev and Kenneth (198la, 1981b, and other papers in
preparation) also accept the karyology of primary spores
and morphology of sporophores as characters with generic
importance, but do not recognize the mechanism of spore
discharge to be taxonomically significant.

No formal proposal of yet another new generic classi-
fication, seems appropriate at this time. This study has
sought only to evaluate the characters and criteria which
have been or might be used to construct generic classifi-
cations of the Entomophthorales. I have extended the con-
clusions of this study regarding appropriate generic cri-
teria to the existing classification schemes in order to
show their net effects and to stimulate further discussion
and debate.

234

Each attempt to propose a new generic classification
for the Entomophthorales has suffered from the effects of
one or more faultily circumscribed genera and from the less
obvious effects of incompleteness. It is clear from the
Still incomplete catalogue of species described as or at-
tributable to Entomophthora (sensu lato) (MacLeod 1963;
MacLeod and Muller-Kogler 1970, 1973; MacLeod et al. 1976),
that the available information about a distressing number
of species is inadequate to allow a reliable classification
in the narrowly defined genera comprising the contemporary
classification schemes. I must also be noted that all of
these latter-day efforts to reclassify the Entomophthorales
have concentrated almost exclusively upon the entomopatho-
genic genera: Several morphologically diverse genera are
known only as saprobes or pathogens of fern gametophytes,
desmide algae, tardigrades, and nematodes; at least three
species of this order are also known to affect humans and
other vertebrates (King 1979). These fungi are no less
valid members of the Entomophthorales than those species
altackinesinSects*+ 1 tSminappropMlatemtOaroUceapOU Gustine
choice of criteria used to define genera or about the cor-
rectness of one or another classification of the Entomoph-
thorales without taking these non-entomogenous genera into
full account.

Despite these impediments to the proposition of a con-
prehensive generic scheme, however, I believe that most
concerned parties now agree that following a period of fur-
thered1scussiony of taxonomi Cucti tenia wel tewit EpempoSssiL le
to propose a generally acceptable, realistic, and phylo-
genetically based classification for the Entomophthorales
in which each genus will have a uniformly high degree of
morphological and biological homogeneity. Despite the
obvious difficulties in proposing such a comprehensive
scheme, some comfort should be found in the thought that IF
the generic scheme which is yet to emerge is accurate, then
no further major adjustments of generic circumscriptions
should be necessary. As new taxa are found and described,
and as older taxa being held in abeyance become better
known, they will be classified in the existing stable gene-
ric structure or seen to differ in enough significant char-
acters to warrant the erection of new genera.

ZOD

ACKNOWLEDGEMENTS

I wish to express my profound gratitude to I. Ben-
ZeMey elem Nem COUCI Rem Geek eCnnetis ahem) a khOrts ands G.
Remaudiere for valuable correspondence and discussion of
the taxonomic and nomenclatural issues raised here, and to
IvBen-Ze'ev Jab. Kramers. DoE. McCabe; and Ra) S-tSoper
FOMMUNCImecn tl calerevicwSeotetneemanuscrEpe.

REFERENCES CITED

AINSWORTH, G.C. 1961. Atnsworth & Bisby's Dictionary of the Fungt,
Sth ed. Commonwealth Mycological Institute, Kew.

AINSWORTH, G.C., F.K. SPARROW, and A.S. SUSSMAN. 1973. The Fungt: An
Advanced Treattse, Vol 4a. Academic Press, New York.

ALEXOPOULOS, C.J., and C.W. MIMS. 1979. Introductory Mycology, 3rd
ed. J. Wiley & Sons, New York.

BALAZY, S. 1978. A peculiar type of the conidia in Entomophthoraceae.
ADStrene LLC COllOdmeInVeLCCOL Ee LAtNO lower acue CZeCNOS.Lang Da >.

BATKO, A. 1964a. Remarks on the genus Entomophthora Fresenius 1856
non Nowakowski 1883 (Phycomycetes: Entomophthoraceae). Bull. Acad.
POLO APOC ome Pe moC he a BLO Lele 8 o19=5218

BATKO, A. 1964b. On the new genera: Zoophthora gen. nov., Trtplospor-
tum (Thaxter) gen. nov. and Entomophaga gen. nov. (Phycomycetes:
Entomophthoraceae). Bull. Acad. Polon. Sei., Sér. Set. Biol. 12:
323-326.

BATKO, A. 1964c. Remarks on the genus Lamta Nowakowski 1883 vs. Cult-
ctcola Nieuwland 1916 (Phycomycetes: Entomophthoraceae). Bull.
Acad. Poton. Set., Sér. Set. Btol. 12: 399-402.

BATKO, A. 1964d. Some new combinations in the fungus family Entomoph-
thoraceae (Phycomycetes). Bull. Acad. Polon. Set., Sér. Set. Btol.
12: 403-406.

BATKO, A. 1964e. Notes on entomophthoraceous fungi in Poland. é£nto-
mophaga, mém. hors sér. 2: 129-131.

BATKO, A. 1966. On the subgenera of the fungus genus Zoophthora Batko
1964 (Entomophthoraceae). Acta Mycol. 2: 15-21.

BATKO, A. 1974. Filogeneza a struktury taksonomiczne Entomophthor-
aceae. In Ewolucja btologicana: Szktce teoretyczne t metodologi-
ezne (C. Nowinskiego, ed.). Polska Akad. Nauk, Inst. Filozof.
Socjol., Wroctaw.

BATKO, A., and J. WEISER. 1965. On the taxonomic position of the fun-
gus discovered by Strong, Wells, and Apple: Strongwellsea castrans
gen. et sp. nov. (Phycomycetes: Entomophthoraceae). J. Invertebr.
Pathol. 7: 455-463.

BENJAMIN, R.K. 1962. A new Bastdtobolus that forms microspores.

ALLSO= ofa 25-2555

236

BEN-ZE'EV, I., and R.G. KENNETH. 1979. JZoophthora erinacea sp. n.
(Zygomycetes: Entomophthoraceae), a fungal parasite of aphids.
Mycovanon 10." 219-252.

BEN-ZE'EV, I., and R.G. KENNETH. 198la. JZoophthora ortentalts sp. nov.,
a fungal pathogen of Aphts cttrtcola (Homoptera: Aphididae), and
two new combinations of other species of Entomophthoraceae. Phyto-
pardstticda 95% 35-42.

BEN-ZE'EV, I., and R.G. KENNETH. 1981b. JZoophthora radtcans and
Zoophthora petchit sp. nov. (Zygomycetes: Entomophthorales), two
species of the 'sphaerosperma group" attacking leaf-hoppers and
frog-hoppers (Homoptera). Entomophaga 26(1): in press.

BEN-ZE'EV, I., and A. UZIEL. 1979. Monellta costalts (Fitch), a new
host for Zoophthora radicans (Brefeld) Batko and Entomophthora
planehontana Cornu (Zygomycetes: Entomophthoraceae) in Israel.
Phiycoparasteted 7: 159-167 -

BERDAN, H. 1938. Revision of the genus Ancylistes. Mycologta 30:
396-415.

BESSEY, E.A. 1950. Morphology and Taxonomy of Fungt. McGraw-Hill
Book Co., New York.

BLACKWELL, M., R.A. SAMSON, and J.W. KIMBROUGH. 1980. Termttartopsts
cavernosa, gen. et sp. nov., a sporodochial fungus ectoparasitic on
ants. Mycotaxon 12: 97-104.

BREFELD, O. 1873. Untersuchungen uber die Entwicklung der Zmpusa mus-
cae und Empusa (Entomophthora) radtcans und die durch sie verur-
sachten Epidemien der Stubenfliegen und Raupen. Abh. Waturf. Ges.
Hol aia loo

BREFELD, 0. 1877. Ueber die Entomophthoreen und ihre Verwandten.

Bot. Zettung (Letpatg) 35: 345-355. 368-372.

BROBYN, P.J., and N. WILDING. 1977. Invasive and developmental pro-
cesses of Entomophthora species infecting aphids. Trans. Brit.
Mycol. Soc. 69: 349-366.

CARNER, G.R. 1976. A description of the life cycle of Entomophthora
sp. in the two-spotted spider mite. J. Invertebr. Pathol. 28:
245-254,

CARNER, G.R., and T.D. CANERDAY. 1968. Field and laboratory investi-
gations with Entomophthora fresenit, a pathogen of Tetranychus spp.
J. Eeon. Entomol. 61: 956-959.

CAVARA, F. 1899a. I nuclei delle Entomophthoreae in ordine alla filo-
genesi-dj queste piante. “Boll. Soc. Bot. ital. 29: 55-60.

CAVARA, F. 1899b. Osservazioni citologiche sulle Entomophthoreae.
Nuov. Giorn. Bot. Ital. (N.S.) 6: 411-466.

COUCH, J.N. 1938. The genus Septobastdium. University of N. Carolina
Presss Guapeleniid-

COUCH, J.N. 1939. A new Contdtobolus with sexual reproduction. Amer.
Je BOCA 26: 119-130.

COUCH, JN. R:V. ANDREEVA, M.° LAIRD; sand’ R°AS NOLANS) 19797) abeano-
myces mtlkot (Dudka and Koval) emended, genus novum, a fungal
pathogen of horseflies. Proce: Nat. Acad: Set. UsS.Av 762 2299-2302.

CUTTER, V.M., Jr. 1942a. Nuclearsbehavior in the Mucorales rier ihe
Mucor pattern. Bull. Torrey Bot. Club 69: 480-508.

CUTTER, V.M., Jr. 1942b. Nuclear behavior in the Mucorales. II. The
Rhizopus, Phycomyces, and Sporodinta patterns. Bull. Torrey Bot.
Club 69: 1592-616.

DAVIDSON, J.G.N., and G.L. BARRON. 1973. Nematophagous fungi: Mertst-
aerum. -Canad. J. Bot. 97° 2314233.

DRECHSLER, C. 1940. Three fungi destructive to free-living terricolous

rato

nematodes. J. Wash. Acad. Sct. 30: 240-254.

DRECHSLER, C. 1951. An entomophthoraceous tardigrade parasite produc-
ing small conidia on propulsive cells in spicate heads. Bull.
torrey Bou. CluDe76: 185-200.

DRECHSLER, C. 1964. An odorous Bastdtobolus often producing conidia
plurally and forming some diclinous sexual apparatus. Amer. d.
DOGO teed iis

DYKSTRA, M.J. 1974. An ultrastructural examination of the structure
of asexual propagules of four mucoralean fungi. Mycologta 66:
477-489.

FITZPATRICK, H.M. 1930. The Lower Fungt: Phycomycetes. McGraw-Hill
Book Co., New York.

GALLAUD, I. 1905. Etudes sur une Entomophthorée saprophytique. Ann.
Ser eNGLUr eon. (desser, 4 da 101=154.

GIARD, A. 1888. Fragments biologiques. XI. Sur quelques Entomophtho-
rées. Bull. Set. France Belgtque 19: 298-309.

GOLDSTEIN, B. 1929. A cytological study of the fungus Massospora
etcadina, parasitic on the 17-year cicada, Magictcada septemdectim.
Amer. J. Bot. 16: 394-401.

GUSTAFSSON, M. 1965. On the species of the genus Entomophthora Fres.
in Sweden. I. Classification and distribution. Lantbrukshogskolans
Ay ole VO S=212%

HALL, I.M., and J.V. BELL. 1963. The synonymy of Empusa thaxtertana
Petch and Entomophthora tgnobtits Hall and Dunn. J. Insect Pathol.
oy 182-186.

HALL, I.M., and P.H. DUNN. 1957. Entomophthorous fungi parasitic on
the spotted alfalfa aphid. Atlgardia 27: 159-181.

HORI, S. 1906. Entomogenous fungi of Japan, 2. Konehi-gaku Zassht
(Bnvomol. Mag., VOoRVO) 5: 81-83.

HUMBER, R.A. 1975. Aspects of the biology of an insect-parasitic fun-
gus, Stronguellsea magna (Zygomycetes: Entomophthorales). Ph.D.
dissertation, University of Washington, Seattle.

HUMBER, R.A. 1976. The systematics of the genus Stronguellsea (Zygo-
mycetes: Entomophthorales). Mycologta 68: 1042-1060.

MUMBER; R.A. 1978. #ntomophthora tgnobtits, the correct name for a
common fungal pathogen of aphids. Mycologta 70: 208-210.

HUMBER, R.A. 198la. Strongwellsea vs. Hrynta: the case for a phylo-
genetic classification in the Entomophthorales (Zygomycetes). In
preparation for Mycotaxon.

HUMBER, R.A. 198l1b. Nuclear cytology and the definitive separation of
Contdtobolus from Entomophthora (sensu lato). In preparation for

. Myecotaxon.

PUMBERGe hie Reo. tOUPRER, ©D-M-.MacLEOD, DD. TYRRELG: R.G KENNETH, and al
BEN-ZE'EV. 1981. Proposal to conserve the entomopathogenic fungal
genus Trtplosporium (Thaxter) Batko against Neozygites Witlaczil
(Entomophthorales). Taxon 30: 187-191. 7

HUMBER, R.A... R-S. SOPER, N. WILDING, and G. REMAUDIERE. 1977. The
identification of certain widely used strains of Entomophthora
pathogenic for aphids. Mycotaxon 5: 307-310.

HUTCHISON, J.A. 1963. The genus Entomophthora in the western hemi-
sphere. Trans. Kansas Acad. Sct. 66: 237-254.

INGOLD, C.T. 1934. The spore discharge mechanism of Bastdtobolus
ranarum. New Phytol. 33: 274-277.

INGOLD, C.T. 1971. Fungal Spores: Thetr Liberation and Dispersal.
Clarendon Press, Oxford.

KELLER, S. 1978. Entomophthora gtgantea sp. nov. and E. caroltintana

238

(Thaxter) comb. nov., two pathogens of Tipula paludosa Meig.
Sydowta Ann. Mycol. 31: 87-93.

KELLER, S. 1980. Two new species of the genus Zoophthora Batko (Zygo-
mycetes, Entomophthoraceae): Z. lanceolata and Z. crassttuntcata.
Sydowta Ann. Mycol. 33: in press.

KENNETH, R.G. 1977. Entomophthora turbtnata sp. n., a fungal parasite
of the peach tree aphid, Pterochlorotdes perstcae (Lachnidae).
Mycotaxon 6: 381-390.

KING, D.S. 1976a. Systematics of Contdtobolus (Entomophthorales) using
numerical taxonomy. I. Biology and cluster analysis. Canad. J. Bot.
of: 45-64.

KING, D.S. 1976b. Systematics of Contdiobolus (Entomophthorales) using
numerical taxonomy. II. Taxonomic considerations. Canad. J. Bot.
04: 1285-1296.

KING, D.S. 1977. Systematics of Contdtobolus (Entomophthorales) using
numerical taxonomy. III. Descriptions of recognized species. Canad.
Jel BOL oo. F1S-129..

KING, D.S. 1979. Systematics of fungi causing entomophthoramycosis.
Mycologta 71: 731-345.

KING, D.S., and R.A. HUMBER. 1981. Identification: Entomophthorales.
In Mtcrobtal Control of Pests and Plant Diseases (H.D. Burges, ed.),
p. 107-127. Academic Press, New York.

KRAMER, J.P. 1980. The house-fly mycosis caused by Entomophthora mus-
cae: Influence of relative humidity on infectivity and conidial
germination. J. New York Entomol. Soc. 88: 236-240.

KRAMER, J.P. 1981. A mycosis of the blood-sucking snipe-fly Symphoro-
myta htrta caused by Erynta tthacensts sp. n. (Entomophthoraceae) .
Mycopathologia: in press.

KRENNER, *J.A- “1961. “Studies in the field of the microscopre fungi.
III. On Entomophthora aphtdis H. Hoffm. with special regard to the.
family of the Entomophthoraceae in general. Acta Bot. Acad. Sct.
HUNG (is) 545-510.

LAKON, G. 1919. Die Insektenfeinde aus der Familie der Entomophtho-
reen. JZ. Angew. Entomol. 5: 161-216.

LATGE; J.-P. 1976.5 Etude morphologique, et cytologique de la formation
de "spores durables" chez une espéce d'Entomophthorale. Compt.
Rend. Hebd. Séances Acad. Sct. 282D: 605-608.

LATGE, J.-P., D.S. KING, and B. PAPIEROK. 1980. Synonymie de Entomoph-
thora virulenta Hall et Dunn et de Contdtobolus thrombotdes
Drechnsler weiWicOvaxcor wal we 32 OO

MacLEOD, D.M. 1956. Notes on the genus Empusa Cohn. Canad. J. Bot.
54: 16-26.

MacLEOD,, D.M. 1963. Entomophthorales infections. In Insect Pathology:
An Advanced Treattse (E.A. Steinhaus, ed.), Vol. 2, p. 189-231.
Academic Press, New York. |

MacLEOD, D.M., and E. MULLER-KOGLER. 1970. Insect pathogens: species
originally described from their resting spores mostly as Tartchtum
species (Entomophthorales: Entomophthoraceae). Mycologta 62: 33-66.

MacLEOD, D.M., and E. MULLER-KOGLER. 1973. Entomogenous fungi: Ento-
mophthora species with pear-shaped to almost spherical conidia
(Entomophthorales: Entomophthoraceae). Mycologta 65: 823-893.

MacLEOD, D.M., E. MULLER-KOGLER, and N. WILDING. 1976. Entomophthora
species with E. muscae-like conidia. Mycologta 68: 1-29.

MAY; Bz, Do WeeROBERTS, and’ Reo, SOPER? “1979"s sintraspecitice genetic
variability in laboratory strains of Entomophthora as determined by
enzyme electrophoresis. Exper. Mycol. 3: 289-297.

239

NEMOTO, H., and A. AOKI. 1975. Entomophthora floridana (Entomophthor-
ales: Entomophthoraceae) attacking the Sugi spider mite, Oltgonychus
hondoensts (Acarina: Tetranychidae), in Japan. Appl. Entomol. Zool.
10: 90-95.

NEMOTO, H., M. KOBAYASHI, and Y. TAKIZAWA. 1979. Scanning electron
microscopy of Entomophthora (Triplosportum) flortdana (Zygomycetes:
Entomophthorales) attacking the Sugi spider mite, Oltgonychus hon-
doensts (Acarina: Tetranychidae). Appl. Entomol. Zool. 14: 376-382.

NOWAKOWSKI, L. 1877. Die Kopulation bei einigen Entomophthoreen.

Bot. Zettung (Letpatg) 35: 217-222.

NOWAKOWSKI, L. 1881. O grupie owadomorkéw (Empusaceae). Dzatenn. III
Zjazdu Lek. Prayr. Polak. Krakow, Seke. Bot. 6: 67.

NOWAKOWSKI, L. 1882. Uber die Entomophthoreen. Bot. Zettung (Letp-
zig) 40: 560-561.

NOWAKOWSKI, L. 1883. Entomophthoreae. Przyczynek do znajomoSci
pasorzytnych gryzbké6w sprawiajacych pomor owadéw. Pamtetn. Wydz.
Akad. Umtiej. w. Krakéw 8: 153-183.

OLIVE, E.W. 1906. Cytological studies on the Entomophthoreae. I. The
morphology and development of Empusa. Bot. Gaz. (Crawfordsville)
41: 192-208.

PAGE, R.M., and R.A. HUMBER. 1973. Phototropism in Contdtobolus coro-
natus. Mycologta 65: 335-354.

PETCH, T. 1938. Notes on entomogenous fungi. Trans. Britt. Mycol. Soc.

21: 34-67.
PETCH, T. 1944. Notes on entomogenous fungi. Trans. Brit. Mycol. Soe.
Chemo = 95s

POHLAD, B.R., and E.C. BERNARD. 1978. A new species of Entomophthor-
ales parasitizing tardigrades. Mycologta 70: 130-139.

REMAUDIERE, G., and G.L. HENNEBERT. 1980. Révision systématique de
Entomophthora aphtdts Hoffm. in Fres. Description de deux nouveaux
pathogénes dtaphidess. Mycotaxon Jie8 269-521.

REMAUDIERE, G., and S. KELLER. 1980. Reconsidération systématique des
genres d'Entomophthoraceae 4 potentialité entomopathogéne.
Mycotaxon 11: 323-338.

REMAUDIERE, G., J.-P. LATGE, and B. PAPIEROK. 1979. Reconsidération
taxonomique de Entomophthora obscura Hall et Dunn. Amn. Mtcrobtol.
(sce nrasteur)mio0: 151-1625

RICHARDSON, M.J. 1970. Ballocephala verrucospora sp. nov., parasitiz-
ing tardigrades. Trans. Brit. Mycol. Soc. 55: 307-309.

RIDDLE, L.W. 1906. On the cytology of the Entomophthoraceae. Proc.
Amer. Acad. Arts Set. 42: 177-197.

ROBERTS, D.W., and R.A. HUMBER. 1981. Entomogenous fungi. In Btology

BeOvaconicdialeringiy(G. 0. ColesandawW.B.skendri ckaweds=).5.V0l.)2,5p-
201-236. Academic Press, New York.

SAMSON, R.A., P.M.J. RAMAKERS, and T. OSWALD. 1979. Entomophthora
thrtpidum, a new fungal pathogen of Thrips tabact. Canad. J. Bot.
DAL OETA S25.

SAWYER, W.H. 1931. Studies on the morphology and development of an
insect-destroying fungus, Entomophthora sphaerosperma. Mycologta
25 Att 432.

SCHROTER, J. 1897. Entomophthorineae. In Die naturlichen Pflanzen-
familten (A. Engler and K. Prantl, eds.), I. Teil, Abt. 1: 134-141.

SELHIME, A.G., and M.H. MUMA. 1966. Biology of Entomophthora floridana
attacking Futetranychus bankst. Fla. Entomol. 49: 161-168.

SHIMAZU, M. 1976. Entomophthora delphacts isolated from the brown
planthopper, Wilaparvata lugens (St41l). Jpn. J. Appl. Entomol.
Zool. 20: 144-150.

240

SHIMAZU, M. 1977. Infectivity of Entomophthora delphacts (Entomoph-
thorales: Entomophthoraceae) to the cotton aphid, Aphts gossypit
(Hemiptera: Aphididae). Appl. Entomol. Zool. 12: 200-201.

SNELL, WoHay and BVAy DICKS 91971, “AlGlossarp er Micology a revised ed.
Harvard University Press, Cambridge.

SOPER, R.S. 1974. The genus Massospora, entomopathogenic for cicadas,
Part I, Taxonomy of the genus. Mycotaxon 1:13-40.

SOPER, R.S. 1981. New cicada pathogens: Massospora ctcadettae from
Australia and Massospora pahartae from Afghanistan. Mycotaxon 18:
50-58.

SRINIVASAN, M.C.,M.J. NARASIMHAN, and M.J. THIRUMALACHAR. 1964. Arti-
ficial culture of Entomophthora muscae and morphological aspects for
the differentiation of the genera Entomophthora and Contdtobolus.
Mycologta 56: 683-691.

STRONG, F.E., K. WELLS, and J.W. APPLE. 1960. An unidentified fungus
parasitic on the seed-corn maggot. J. Heon. Entomol. 53: 478-479.

THAXTER, R. 1888. The Entomophthoreae of the United States. Mem.
Boston Soe. Natur. Htst. 4: 133-201.

THAXTER, R. 1920. Second note on certain peculiar fungus-parasites of
living insects. Bot. Gaz. (Crawfordsville) 69: 1-27.

TSINTSADZE, K.V., and S.G. VARTAPETOV. 1976. A new fungus, Entomoph-
thora adjartca sp. n. (Phycomycetes, Entomophthoraceae), affecting
Tetranychus urttcae Koch. Bull. Acad. Set. Georgian SSR 83: 465-468.

TURIAN, G. 1957. Entomo-mycoses dans la région de Genéve. Mtt.
Schwetz. Entomol. Ges. 30: 93-98.

TYRRELL, D., and D.M. MacLEOD. 1972. A taxonomic proposal regarding
Delacrotxta coronata (Entomophthoraceae). J. Invertebr. Pathol. 20:
PPar5 5

VUILLEMIN, P. 1886. Etudes biologiques sur les champignons (Entomoph-
thora gloeospora Vuill.). Bull. Soe. Set. Nancy 8: 34-46.

VUILLEMIN, P. 1895. Deuxiéme notice sur les travaux scientifiques.
Bull. Soe. Set. Naney (cited by Cavara 1899a-b).

VUILLEMIN, P. 1900a. Développement des azygospores chez les Entomoph-
thorées. Compt. Rend. Hebd. Séances Acad. Sct. 130: 522-524.

VUILLEMIN, P. 1900b. Développement des azygospores chez les Entomoph-
thorées. Compt. Rend. Assoc. Frang. Avancem. Set. 180: 670-685.

WATERHOUSE, G.M. 1973. Entomophthorales. In The Fungt: An Advanced
Treatise (G.S. Ainsworth, F.S. Sparrow, and A.S. Sussman, eds.),
Vols 4b% p> (219-229 “vAcademicePress , New ork.

WATERHOUSE, G.M. 1975. Key to the species of Entomophthora Fres.
BULED EYL. MY COL poOC moe wl O—4 1

WILDING, N. 1970. Resting spore formation and germination in &. fre-
Seni. “Rothamsted fap wy Sta. Rpt. for 1970, 12t. 1°) 270.

WITLACZIL, E. 1885. WNeozygttes aphtdis, eine neue Gregarine. Arch.
Mtkrosc. Anat. 24: 599-603.

ZIMMERMANN, G. 1978. Zur Biologie, Untersuchungsmethodik und Bestim-
mung von Entomophthoraceen (Phycomycetes: Entomophthorales) an
Blattlausen. Z. Angew. Entomol. 88: 241-252.

——————

MYCOTAXON

VO lee’ CULeNO mls) DD.w24.1-256 April-June 1981

LES HYMENOCHAETE A ELEMENTS HYMENIENS PINNATIFIDES

DR CemLEGEN

Labonatoine de Mycologie assocté au C.N.R.S.
Université Claude Bernartd-Lyon I . Bat. 405.
43, Bd du 11 Novembre 1918. F-69622 VILLEURBANNE

SUMMARY

Hymenochaete acanthophysata nov.sp. with acanthophysate hy-
phae and Hymenochaete hauerslevti nov.sp. with pseudoacantho-
physes are described. The author compares these African spe-
cies with the only two known species with pinnatifid-tipped
hyphidia in their hymenium: H. digitata Burt and H. pinnati-
ida Burt. The later has been found again: descriptions of a
non stratose specimen from Africa and of a two-layered spe-
cimen from Guadalupe are given. An abstract in English inclu-
ding a detailed key to species is given in the end of the pa-
per.

INTRODUCTION

Dans le cadre d'une étude des spécimens du genre Hymeno-
chaete récoltés principalement en Afrique par J. BOIDIN d'une
part et G. GILLES d'autre part, nous avons décrit précédem-
ment une espéce, H. spathulata, remarquable par ses spinules
a sommet spatulé (LEGER, 1980) ainsi que deux espéces, H. 4e-
panabihis et H. hatpago (LEGER, 1981) a spinules pourvues de
diverticules.

Le présent travail a trait aux espéces dont 1'hyménium
posséde des éléments stériles ornés de digitations. Jusqu'ici
deux espéces présentent ce caractére: H. pinnatifida Burt et
H, digitata Burt. H. pinnatifida a été retrouvé en République
Centrafricaine, au Gabon et a la Guadeloupe. Une description
compléte est donnée qui fait apparaitre notamment que cette
espéce peut étre bistratifiée. A titre comparatif, l'étude de
He OLO. 01d estereprisesebecompletce aupartim dustypesaesnUR..
Enfin, deux espéces sont décrites : H. acanthophysata nov.sp.
a hyphes acanthophysoides et H. hauetslevii rov.sp. a éléments
hyméniens de type pseudoacanthophyses.

242

HYMENOCHAETE PINNATIFIDA Burt
ANN ee MESSOULT RbOU. sGard wnt ro Do eam iO

Basidiome résupiné, étalé, adhérent, mince, 60-150-
(200) pm, en petites taches plus ou moins confluentes, brun
jaunatre (Munsell 1OYR 5/4 ou encore Saccardo's umber selon
Ridgway) puis brun rouillé (Munsell 5YR 4/6) quand il s'éta-
le plus largement, 4 marge fibrilleuse appliquée, 0,5 mn,
chamois (Munsell l1OYR 7/8).

Cortex (Cx, fig.1) épais de 5 4 15 um, sombre, formé
d'un enchevétrement serré d'hyphes brun-rouge foncé, plus ou
moins cimentées et collapsées, x 2-2,5-(3) pm, 4 paroi ép-
aisse.

Contexte (C, fie) assez elache, 50 a5 100simadsepaise
seur, dimitique. Hyphes génératrices hyalines, x 1,5-2 pm,

a paroi mince, ramifiées et cloisonnées. Hyphes squeletti-
ques brun-jaunatre, x 2-2,5 pm, a paroi épaisse, non cloi-
sonnées, exceptionnellement ramifiées.

SpinwLessbrun) Ponce.) laspLupactal on j0mues 5. mum.
non émergentes; certaines, plus rares, mesurent 40-50 x 6-

7 wm et émergent jusqu'a 20 pm. Toutes ces spinules nais-
sent d'une couche sous-hyméniale sombre.

Hyménium (H, fig.1) de 15 a 20 pm; rares basidioles
et trés rares basides mures, 15 x 3 pm, a 4 stérigmates de
3 pm. Dendrophyses jaunes trés abondantes, longues de 12
a 22 pm et de diamétre variant de 2 4 6 pm, a4 nombreuses
ramifications tres etroites (O25 pm) en petites touffes
terminales pour la plupart. Trés rares hyphes paraphysoides
gréles grimpant le long de certaines grandes spinules. Assez
nombreuses masses cristallines volumineuses (souvent de 10-
20 x 10 pm) a4 la base de 1'hyménium.

Sous-hyménium (sH, fig.1) formant une zone sombre de
20 pm environ, d'aspect identique au cortex, constituée d'
hyphes plus ou moins cimentées et collapsées, x 2-3 pm, 4
paroi épaisse.

Spores (sur sporée) cylindriques étroites, trés légére-
ment déprimées, 3,8-4,5-(5) x 1,6-2 pm, hyalines, uninucléees
(Giemsa), non amyloides, blanches en masse.

Description effectuée 4 partir de la récolte LY 5584, La Ma-
boké, République Centrafricaine, 31 mars 1965, leg. J. Boi-
din (Herb. Boidin).

AUTRES RECOLTES
LY 5931 sur Carapa procera, La Maboké, République Centra-
fricaine, 16 septembre 1967, leg. J. Boidin.

243

(a>
¢

oP -

Gans iy
One

Or
aaah

a
Bo.

Se
Gis

Hymenochaete pinnatifida Burt
(LY 5584 )

244

LY 6041 sur support indéterminé, Boubakiti, République
Centrafricaine, 27 septembre 1967, leg. J. Boidin.

LY 9260 sur support indéterminé, Libreville, Gabon, 20
novembre 1978, leg. G. Gilles.

ESPECES DE REFERENCES EXAMINEES

H. pinnatifida Burt, New Smyrna, Florida, january 1897,
CrCl iloydee SUeGlype) pete 2140. ICED rn

H. pinnatifida Burt, ad Ligna emortua angtospernae in
paroecta Aurelia, Algiers, 17 september 1971, leg. M.T.
Dunn, det. A.L. Welden (Fungi Ludovictant ex herbarto Tula-
neav/ Gb5) eevee oe

LE: GAS PARTECULIER* DE LA RECOLTE EY%s091]

Ce spécimen, trouvé par J. BOIDIN le 29 septembre 1976 4
la Guadeloupe se présente comme un H. pinnatifida bistra-
ti BLes Chien).

Tout se passe comme si un deuxiéme basidiome complet avec
un cortex Cx2, un contexte C2, une zone sous-hyméniale sH2
et un hyménium H2 (pratiquement stérile) etait né d'un ba-
sidiome précédent constitué (comme décrit pour LY 5584) d'
un hyménium Hl], d'un sous-hyménium sHl, d'un contexte Cl et
d'un cortex basal Cxl. Dans le dessin de la figure 2, la ba-
se du champignon —- c'est-d-dire le cortex Cxl ainsi que les
4/5 du contexte Cl - n'est pas représentée. Le basidiome a.
une €paisseur totale de 150 a 230 pm dont 100 pm pour le
contexte Cl (alors que le contexte C2 n'a qu'une quinzaine
de pm d'épaisseur).

DISCUSSION :

Les spécimens africains présentent une trés bonne concor-
dance avec le type d' H. pinnatif{ida Burt examiné ainsi qu'
avec la récolte américaine LY 6723 déterminée par A.L. WEL-
DEN. Cependant, deux remarques s'imposent

- L'assez grande variabilité de couleur des basidiomes,
non seulement d'une récolte a4 l'autre mais aussi a4 l'inté-
rieur d'une méme récolte. Nous distinguons trois groupes
selon la couleur: gris rosatre (light cinnamon drab, vers 2,5
YR 6/2) pour LY 9260 et pour le centre de quelques taches de
LY 6/233) brun; rougeatre (reddish brown, vers: sv bu) se ae le
ou 4/6) pour LY 6041, LY 8091, LY 6723 pro parte et LY 5584
pro parte; brun jaunatre (yellowish brown, vers l10YR 5/4)
pour LY 5584 pro parte (petites taches trés jeunes) et LY
5931. Cette couleur variable s'explique au moins en partie
par le développement relatif de 1l'hyménium: ainsi, les taches
de LY 6723 ot 1'hyménium est bien développé et fertile sont
gris rosatre alors que les taches brun rougeatre ne présen-

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ae Way
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Eee oy
‘Saas
ae
se . Bin. ae
2) “ .
Aes a
SITs iy x
3: ae Ue
A. i
/ A
a i
Fl
AQ

Figure 2: partie supérieure d Hymeno-
chaete pinnatifida Burt (LY 8091)

246

-tent qu'un hyménium faiblement développé, pratiquement sté-
rile et constitué presque exclusivement de dendrophyses. De
méme LY 9260, dont 1l'hyménium fertile est bien développé est
uniformément gris rosatre. La marge, chamois a rouillé, a
presque toujours une couleur plus vive que le reste de 1"hy-
ménium.

- Concernant la stratification, 1l'examen de la récolte
LY 8091 conduit incontestablement 4H. pinnatifida Burt.
Cette bistratification n'a jamais été signalée ni par E.A.
BURT, ni ultérieurement lors d'autres récoltes.

En résumé, H. ptnnati4ida Burt est une espéce bien carac-
térisée par ses dendrophyses (a nombreuses et fines ramifi-
cations)issues d'une zone sous-hyméniale sombre, par un
contexte dimitique assez lache et un cortex basal sombre. La
couleur de l'hyménium peut varier de facon sensible suivant
1'état de maturité tandis que la marge est toujours de cou-
plus vive.

L'aire de dispersion de H. pinnatif{ida s'étend non seulement
a l'Afrique mais également 4 la Guadeloupe dont le spécimen
montre un caractére jamais observé jusqu'ici pour cette espé-
ce: une bistratification nette. Il serait intéressant de re-
chercher aux Etats-Unis notamment, une présence éventuelle

de stratification lors de futures recoltes.

HYMENOCHAETE DIGITATA Burt
Ann Missouri Bot a) Gards>= O47 OLS

Basidiome résupiné, largement étalé, adhérent, épais
de 700 a 800 pm, brun havane (Brussels brown 4 antique bro-
wn, 7,5YR 4/4 a 5/6), 4 surface granulée-tuberculée, 4 marge
trés mince, évanescente, concolore.

Cortex (Cxl, fig.4) d'une quinzaine de pm, formé d'un
enchevétrement serré et brun rouge sombre d'hyphes de 2,5-3
pm de diamétre, plus ou moins cimentées et collapsées, 4 pa-
rol épaisse.

Ce cortex se prolonge par un feutrage assez lache constituant
le revétement piléique (tomentum T), d'environ 40 ym d'épais-
seur.

Contexte (fig. 3 et 4) stratifié, composé d'une alter-
nance d'une quinzaine de couches sombres avec un nombre égal
de couches plus claires. Les couches sombres (appelées Cx2 4a
Cx8 et dont seules Cx7 et Cx8 sont représentées sur la figure
3) ont une teinte brun rouge foncé et présentent une consti-
tution identique 4 celle du cortex basal Cxl. Les couches
plus claires ont une double constitution: d'une part, les

Ex
i=

EE ISA SY Be TR) EE qe
NAHB) \ LV Be 4 1 AB
r Guy Riek #0 ee os an.
eg eS at hg se &
G Cl wee soca 2
Ly 4 ; ass Rt Posie AID os O
( ee | PY t) 4H po ek T/
UO ALD
§ pa \)
NY LOE
UAE eH
\G Q Nad LS
WAT \
4a

(VY
aaulnassp
LR aed

Cx7

Figure 3: partie supérieure d’
Hymenochaete digitata Burt (Type)

248

Figure 4: base d* Hymenochaete
digitata Burt (Type)

hyméniums antérieurs (nommés Hl] a H7 et dont seuls H1,H/,H8
sont représentés sur les figures 3 et 4) sont bruns, formés
d'une dense palissade d'hyphes de 2-3 pm de diamétre, 4 pa-
roi épaissie, a articles relativement courts (7 4 10 pm), trés
collapsées, auxquelles se mélent des dendrophyses 4 paroi
épaissie, entiérement jaun&tres. D'autre part, au dessus de
chaque hyménium, des zones trés claires sont formées d'un
enchevétrement trés lache d'hyphes cloisonnées et ramifiées,
de 2-3 pm de diamétre, a4 paroi mince et d'hyphes non cloi-
sonnées, 2-3 pm de diamétre, a paroi épaissie (dimitisme).
A la base de chaque hyménium, de nombreuses masses cristal-
lines plus ou moins volumineuses.

Spinules nées pour la plupart des couches trés som-
bres; elles sont de deux types: trés nombreuses petites

249

spinules 20-30 x 2-3 pm, non émergentes. et grandes spinules
50-75 x 6-7 pm, moins abondantes et émergentes jusqu'a 40 pm.

Hyménium (H8, fig.3) de 25 pm environ; quelques basi-
dioles mais basides extrémement rares (non vues par BURT),
[jexe2.5— et wa 4epetitsesteriematesades!)5enm.) Leessentiel
est constitué par des sortes de pseudoacanthophyses nombreu-
ses: ce sont des éléments hauts de 12 a 17 pm, 4a base ren-
flée (3,5-4,5 pm) hyaline et a paroi mince, tandis que 1'ex-
trémité rétrécie, jaunatre et 4 paroi épaisse porte de nom-
breuses ramifications en tous sens. Certaines sont cependant
déja entiérement jaunatres et a4 paroi épaissie comme celles
des hyméniums antérieurs.

Spores non observées.

Description d'aprés le type de H. digitata Burt, El Boquete,
Chiridut.ePananave Won seMaxOn «moo eC PH

DISCUSSION :

L'étude du type de BURT a permis d'apporter quelques pré-
cisions sur cette espéce qui n'a pas fait l'objet,depuis le
travail originel, d'étude détaillée. Le point le plus inté-
ressant, outre la présence non encore signalée d'un tomentum
et la constitution précise des différentes couches de strati-
fication, est sans doute l'aspect détaillé des pseudoacantho-
physes qui n'avaient été que superficiellement décrites par
BURT.

HYMENOCHAETE ACANTHOPHYSATA Léger nov.sp.

Basidioma resupinatum, Late jacens, adherens, tenue, obscure
braunneum, magis tn medio propter prutnam grtiseokum; ambi tu
ex nrubiginosos fulvo, margine angustissimo, albo; trama bir-
Strata, dimtica, hyphis genetricibus hyalinis, tunica tenut,
x 2-3 pm, ramosis fibulatisque, ex hyphis skeletticis e fla-
vis brunnets, tunica crassa, xX 2-3 pum, neque rAamosis neque
Septatts constante; contice nullo; spinukis braunnets, amplrs,
40-70 x #-8,5 pm, usque ad 35 pum omergentibus ; hyphis para-
physoridets gnacitibus, x 1-1,5 pm, quasdam spinulas ctrcun-
dantibus; basidiis narissimi(s, 12 x 3 pum, 4 sterigmatis 3 pum
Longis; peramultis hyphis acanthophysoridets, tunica crassa,

x 2-2,5 pm, non vel vix emergentibus; passin crystallis in-
ter hymenium praesentibus; sports cylindratis, Leviter de-
pressis, 4-4,5-(5) x 1,5-1,8 pm, hyalinis, uninucleatits,
haud amyloiders, in massa akbrs.

HOLOLU USE: ely on (LY), legends sBoidinen« LY //41% Ini suostia-
to 4ncognito, Makokou, Gabon.

250

Ethymologie : espéce ainsi nommée du fait qu'elle posséde
de nombreuses hyphes acanthophysoides (acanthophyses défi-
Miles) pareAeee IEALTTenml 920).

DESCRIPTION DU TYPE :

Basidiome résupiné, largement étalé, adhérent, mince,
125 a 200 pm, brun sombre (5YR 4/2, fulbtgtneus de Saccardo)
puis plus grisatre au centre par la pruine (5YR 5/2, benzo
brown R. 4 2,5YR 5/2, cinnamon drab R.). Bordure fauve rouil-
1é (7,5YR 5 a 6/6, vers antique brown R.). Marge trés étroi-
te, fibrilleuse, trés appliquée, blanche.

Contexte (fig.5) bistratifié, dimitique. Hyphes géné-
ratrices hyalines, a paroi mince, 2-3 um de diamétre, rami-
fiées et cloisonnées. Hyphes squelettiques brun jaunatre, 4
paroi épaisse, 2-3 pm de diamétre, ni ramifiées ni cloison-
nées.

A la base du champignon (pas de cortex), le contexte est en-
chevétré peu sombre (30-50 pm) puis passe 4 une zone sous-—
hyméniale plus obscure (8-15 jim) ot les hyphes sont plus
fermement enchevétrées et plus ou moins cimentées. Enfin,
les hyphes se redressent pour donner 1'hyménium supérieur.

Spinules brunes, massives, a extrémité peu pointue
mais assez souvent mucronée, 40-70 x 7-8,5 jam, émergentes
jusqu'ad 35 pm. Des hyphes paraphysoides gréles (1-1,5 pm)
peuvent les entourer.

Hyménium : l'ancien, collapsé, comprend des basidioles
fripées, longues de 6-10 pm, des spinules et des éléments
acanthophysoides brun jaunatre. Ces éléments naissent de la
zone sous-hyméniale sombre. L'hyménium supérieur est formé de
basidioles de 6 4 10 (15) pm de haut, de spinules, de trés
rares basides 12 x 3 pm, 4 4 stérigmates de 3 pm et surtout
d'abondantes hyphes acanthophysoides 4 paroi épaissie, de 2 a
2,5 pm de diamétre, non ou peu émergentes (10 pm). Ces sortes
d'acanthophyses a paroi mince lorsqu'elles sont jeunes, nais-—
sent d'hyphes squelettiques ou génératrices. Localement, des
masses cristallines se rencontrent au niveau de 1'hyménium.

Spores (sur sporée) cylindriques, légérement déprimées
4-4,5-(5) x 1,5-1,8 pm, hyalines, uninucléées (Giemsa), non
amyloides, blanches en masse.

Sur support indéterminé, Makokou, Gabon, 5 mai 1976, leg. J.
Boldin. iHolotype »:) LY 9/741 (Herb. Boldin).

AUTRES RECOLTES

Cote d'Ivoire : LY 7053, sur support indéterminé, Abidjan,
5 novembre 1972, leg. G. Gilles.

Gabon : Makokou (leg. J. Boidin), sur support indéterminé
LY 7734, 4emairel 9/63) LY 27 /SO8eteL Ye i/o on wDemat who 70 atic

Poel

anthophysata

ei tears rtinie ere OTE — Ry SE ae \ "
ee — = = = i Y y i: 2) = nee 5 CA aie, . iat
; Ree eee SEE EE eee | pS we a L., y iS = £2) LA Ce

Figure 5: Hymenochaete ac

ies

Klainedoxa gabonensis : LY 7759, 7 mai 1976; sur Anctstro-
VNU CLUES eee Vey OS Osan Sema tenho / O-

Gabon "Libreville (leg! G. Crlles)jj "sur supporteinacter=
mines LY 9114.32 janvier 19/93eLY 915 eaetevrtermi7o oY
919854 mars 1979s LYe92 155 .12imars 1979s LYe925SezOenoven—
bre 1978s" bYe92612 tl Omnovembrem | 97 Oslo 4/0) oma vet ee oe
LY 9298. 22eavril NOD LY 9310. 2S ravines! Oo ee Limo ae.

Amey une OO

DISCUSSION
Ces nombreuses récoltes permettent de mieux cerner les

caractéres de H. acanthophysata, 4 travers les variations de
la couleur de l"hyménium et surtout de la stratification ou
non des é@échantillons

Couleur de 1l'hyménium: l'abondance des hyphes acantho-
physoides, lorsque 1"hyménium n'est que trés peu ou pas fer-
tile, provoque un net assombrissement de la couleur. Ainsi,
LY 7053, LY 7830 et LY 9417 par exemple qui sont a peu prés
stériles ont une couleur brun rougeatre foncé (5YR 4/3 a 3/3
et méme 2,5YR 2/4 pour LY 9298); cette couleur est renforcée
par un aspect velouté souvent trés marqué. Lorsque 1'hyménium
est bien développé par contre, la couleur est plus pale: soit
plus ¢risatres (25 5YRe 5/2. ae4)/ 2senava Pap rowneR sepOU TE YEO Ol
par exemple) soit plus jaunatre (7,5YR 5 a 6/4, wood brown
R. pour LY 7755). Une méme récolte montre assez souvent une
telle variation de couleur: par exemple, LY 7750 est tantot
chocolat (5YR-3/4)cet=veloute,stancolsenissmosatrer (oYRmo 25
benzo brown R.) et aride. De facgon similaire, LY 7755 (ainsi
que 7/759) offre un trés bel exemple: le centre est en effet
gris légérement rosé (5YR 6/2) alors que le reste de la sur-
face est bai ferrugineux (5YR 4/6).

Stratitication:ssurniunatocal deml/,recoltes siscilesm>
présentent une stratificatwon: LY /734,eLY9258 ely LY. 7/741)
(type) sont bistratifiés. LY 9273 a quelques strates mais
seul LY 7750 (dont 1'épaisseur atteint 700 pm) montre 6 4 7
hyméniums successifs. Tous les autres spécimens (épais de 30
4 100 pm) n'offrent aucune stratification. Le basidiome a
dans ce cas une structure plus simple: 4 la base, un mince
contexte assez sombre (15-20 jim) formé d'hyphes enchevétrées
en tous sens donne naissance 4 un contexte vertical dont les
hyphes se terminent presque toutes en éléments acanthophy-
soides; ceux-ci, comme les spinules, sont répartis dans tou-
te l'épaisseur du basidiome. Qu'il s'agisse de spécimens
stratifiés ou non, la plupart des récoltes sont presque ou
totalement stériles: plusieurs sporées n'ont pu étre obte-
nues et les autres n'ont que peu de spores.

En conclusion, H. acanthophysata nov.sp. est une espéce

Zoo

stratifiée ou non, dont le basidiome étalé-adhérent est de
couleur brun foncé, parfois velouté, a4 1'état substérile et
plus grisatre et aride lorsque 1l'hyménium est normalement
constitué (les basides étant de toute facon trés dispersées).
La présence de nombreuses hyphes acanthophysoides est sans
conteste le point le plus remarquable avec les spinules tra-
pues et massives.

HYMENOCHAETE HAUERSLEVII Léger nov.sp.

Basidioma resupinatum, primum tenutssimun, e Luteo ochtaceo,
dein paulo cnassius, cinnamomeum, srustulosum, Laxe ad subs-
cukum e badio ferruginosum adnatum; ambitu indeterminato,
concolone; trama Laxe interamixta, ex hyphis braunnets, tunica
Ancnassata, x 4-5-(6) pum, septatis ramosisque constante, at-
que ex hyphis hyalinis, tunica tenut, x 2-3 pm, septatis
namosisque; cortice nuklo; crystaklis permultis ad et inter
hyphas; spinulis braunnets, acutis, Longis angustisque, 90-
170-(190) x (4)-6-8 pum, usque ad 100 pm emergentibus; hyphis
paraphysoidets gracilibus, x 2 um, spinulas scandentibus ;s
basidtis saepius paulum constrictis, 12-17-(19) x 4-5 pm,

4 asterigmatis 5 pm Longis; pseudoacanthophysibus tunica te-
nut vek Leviter incerassata, non vel vix emergentibus, 10-
13-(20) x 3-5 pum, paucts apicalibus diverticulis 2-3 pm Lon-
gis praeditis; sports cylindratis (6)-6,5-7,5-(8) x 2,8-3-
(3,2) pum, apieulo saepe manifesto, hyalinis, haud amyloiders.

HOCOAUMUAM: aCopenhague’(C)e leo. K. Bjdrnekaer nn, Afr.22, <n
Substrate tneognito, 4n monte Kenya, Kenya, 31-III-1963.

Ethymologie: espéce dédiée au Dr. K. HAUERSLEV qui nous a si-
gnalé et communiqué le spécimen.

DEOCRLPTEONSDUL TYEE

Basidiome résupiné, étalé, d'abord trés mince (60 pm),
ocre jaune (IOYR 6/6, brownish yellow) puis un peu plus épais
(100-110 pm), cannelle (7,5YR 6/7, cinnamon), frustuleux,
Pau ssanteapparaucre un feutrage bar terrugineuxe (oYR 4/6, cred—
dish brown). Marge indéterminée, concolore.

Contexte (fig.6) lachement enchevétré d'hyphes brunes
a paroi @paisse, 4-5-(6) pm de diamétre, cloisonnées et rami-
fiées et d'hyphes hyalines 4 paroi mince, de 2 4 3 pm de dia-
métre, @galement ramifiées et cloisonnées. Nombreux cristaux
sur et entre les hyphes. Pas de cortex basal.

Spinules brunes aigties, étroites et trés longues, 90-
170-(190) x (4)-6-8 pm, naissant a différentes hauteurs,
émergentes jusqu'a 100 pm. Des hyphes hyalines et gréles de

254

2 pm de diamétre grimpent souvent le long de ces spinules.

Hyménium formé de basidioles, de basides assez sou-
vent un peu constrictées, 12-17-(19) x 4-5 jm 4 4 stérigma-
tes de 5 pm et de pseudoacanthophyses 4 paroi mince ou légé-
rement épaissie, 10-13-(20) x 3-5 pm, avec quelques diverti-
cules terminaux (4 4 6 le plus souvent) de 2 a 3 pm de long.
Basides et pseudoacanthophyses sont abondantes dans la par-
tie jeune, ocre jaune, du basidiome et beaucoup plus rares
dans la partie agée, cannelle.

Spores (observées sur les coupes car sporée non obte-
nue ecyluindriquesnm (0) 0.07 on (Oe Xe. Oro (Ole aiee 0 le
cule souvent bien marqué, hyalines, non amyloides.

Sur support indéterminé, Mt Kenya Forest, Kenya, 31 mars
1963, leg. Ke" Bjgrnekaer.-Holotypes: Afr.22.(C).. = petits1s0—
type adresse par K. HAUERSLEV 4° J<)BOLDIN@ens1973 5 GYe7250)<

DISCUSSION

Hymenochaete hauerslevii nov.sp. est un champignon par-
faitement caractérisé par ses pseudoacanthophyses qui rap-
pellent celles de Stereum australe Lloyd (voir J. BOIDIN,
1960, p. 67). Aucune espéce d'Hymenochaete possédant de tels
éléments hyméniens n'a été décrite précédemment. Un autre
caractére intéressant est la grande ressemblance de cette
espéce nouvelle avec H. cinnamomea (Pers.)Bres. : tout d'
abord la couleur cannelle de la surface mais surtout la
structure lorsque H. cinnamomea est jeune et a l'état uni-
stratifié, c'est-a-dire le contexte lache, les spinules lon-
gues, @troites et trés émergentes; méme la taille des spores
rapproche les deux espéces. Cependant, aucune des récoltes
européennes de H. ctinnamomea unistratifié que nous avons pu
examiner ne présente de pseudoacanthophyses, ce qui permet
de distinguer aisémment H. hauerslevit. I1 serait intéres-
sant de rechercher si des spécimens avec pseudoacanthophy-
ses ne se cachent pas parmi les récoltes américaines uni-
stratifiées de H. ctnnamomea.

ABSTRACT

In this paper, species of Hymenochaete were studied which
possess hymenial hyphidia with "pinnatifid tips' according
to BURT's terminology. To the known species with such a fea-
ture, H. pinnatifida Burt and H. digitata Burt, two species
are added : H. acanthophysata nov.sp. and H. hauerslevit nov.
sp.. Although these species seem to form a group of their
own in our study, it would certainly be excessive to consi-
der them as a natural grouping. We give below a diagnostic
key which summarizes the main characters of each species

ji |
a; A
: 6, (Sates
( el \\ / WY oe ow
s, CS (as iy
V4 | cn:
ss y I
Hi fl 4
\/

256

1- Cortex present (Cuticle Sensu REEVES and WELDEN, 1967)

be COLEOX ADSOME wc ricoh sie lou ca cieic cs tec erersec ect n eLneeen a cnn ieee creas 3

2- No tomentum (abhymenial hairs); dark subhymenium of
tightly interwoven hyphae, context not stratose ( but
a two-layered stage observed in LY 8091 from Guadalupe)
and dendrophyses. West Indies, Gabon, Central African
Republic andeGuadaltipem teen H. pinnatifida Burt.

2- Tomentum present; no dark subhymenium; context stratose
of several hymenial layers and particular pseudoacantho-
physes with dendrophyse-like apicis. Panama ...........

Be es eee H. digitata Burt.

3- Context not stratose of loosely interwoven hyphae; slen-
der setae 90-170-(190) x (4)-6-8 pm and pseudoacanthophy-
ses hyaline; sporese(6)-6,0-/,5=(8) x92, 0-5—( on 4) em.
Kenya estes 6 veto eis H. hnauersleur.% Léserii nov. sp,

3-— Context stratose (but found frequently in first-—stratum
stage); setae somewhat massive, 40-70 x 7-8,5 pm, and
very numerous acanthophysate hyphae; spores 4-4,5-(5) x
1,5-1,8 pm. Ivory Coast and Gabon ...-....eeeee wesc rece

Beth oA bee H. acanthophysata Léger nov. sp.

ACKNOWLEDGEMENTS

Thanks are due to H. ROMAGNESI for the translation of La-
tin diagnoses and to Dr. K. HAUERSLEV for sending the speci-
men from Mt Kenya. We are also very grateful to Dr. J. ERIKS-
SON, and Dr. A.U. WELDEN"for critical review wand sto the Curatay
of the Farlow Herbarium for the loan of specimens.

REFERENCES

BOIDIN J. 1960. Le genre Stereum Pers. S.L. au Congo Belge.
Bad ee ard.s Doth Cat eeDrUxe lec. m0 memsn 277 on

BURT E.A. 1918. Thelephoraceae of North America X. Ann. Mis-
Syolbpes, efolun (Cemetsls “Sy 4 Sloss i.

LEGER J.C. 1980. Hymenochaete spathulata nov.sp. (Basidio-
mycétes aphyllophorales). Bull.Soc.Mycol.France,96:407-411.

LEGER J.C. 1981. Un curieux groupe d'Hymenochaete a spinules
denticulées (Basidiomycétes aphyllophorales). Bull. Soc.
Mycol. France (sous presse).

PILAT A. 1926. Monographie der mitteleuropaischen Aleurodis-—
cineen. Ann. Mycol. 24 : 203-230.

REEVES F., Jr. and WELDEN A.L. 1967. West Indian species of
Hymenochaete. Mycologia, 59 : 1034-1049.

MYCOTAXON

WOOL eNO elem) 20 /—250 April-June 1981

CHLORIDIUM AND SOME OTHER DEMATTACEOUS HYPHOMYCETES
GROWING ON DECAYING WOOD. CORRECTIONS AND ADDITIONS

W. GAMS and V. HOLUBOVA-JECHOVA

Centraalbureau voor Schitmmelcultures, Baarn, and
Botantcal Instttute, Czechosl. Academy of Setences,
Pruhontce near Praha.

Soon after publication of our paper under this title
(tn Stud. Mycol. 13. 1976) our attention was drawn to some
homonymies (Index of Fungi 4: 469. 1978) and incomplete-
nesses which we wish to correct here.

CHAETOSPHAERIA FUSISPORA W.Gams & Hol.-Jech., Z.c. p. 45
is a homonym of
Chaetosphaeria fustspora (Kawamura) Hino tm Bull. Miyazaki
Coll. Agric. For. 4: 191. 1932 (Syn. Miyoshta fustspora
Kawamura; Mtyoshtella fustspora (Kawamura) Kawamura) ;
@ possibly congeneric fungus with much wider ascospores,
Reals Se =< lo) ain = and
Chaetosphaerta fustspora P. Larsen tn Dansk bot. Ark. 14(7):
7. 1952; which probably belongs to Chaetosphaerella
Dy Wiel thm Jstevernay

We Cherefore rename our fungus

Chaetosphaerta fustformts W.Gams & Hol.-Jech. nom. nov.
replaced synonym: Chaetosphaerta fustspora W.Gams &
Hole ech mastud savy col. 5-15 aa 1916.

The genus CYLINDROTRICHUM (l.c., p. 48) was incompletely
treated.
Species published before our paper include:
Cylindrotrtchum proltferum Matsushima, Icones Microfungorum
a Matshushima lectorum: 47. 1975,
Cylindrotrichum trtseptatum Matsushima, Z7.c.: 48. 1975 and
Cylindrotrtchum trtseptatum M.B.Ellis, More dematiaceous
Hyphomycetes: 470. 1976 (a homonym) = C. elltstt Morgan-
Jones tn Mycotaxon 5: 490. 1977.

Since our publication four further species were described:
Cylindrotrtchum oblongtsporum Morgan-Jones tn Mycotaxon
5: 487. 1977, which should be compared with our

298

C. 2tgnoéllae,

Cylindrotrtchum gortt Lunghini in Micol. ital. 8: 25. 1979,
distinet from the previously known species by smaller,
often slightly curved conidia, 5.7-9.9 x 1.9-3.0 um,
Cylindrotrtchum heltset forme Marvanova tn Trans. Br. mycol.
Soc 75.) 300 nN 1O TO. wibhs uri cuspidate apical cells of the
conidia and

Cylindrotrtchum curvatum Morgan—Jones in Mycotaxon 12:

250. 1980, with curved conidia, Horus oe SSI NO) Hie,

MYCOTAXON

NOlae eles NO clea) cod -200 AD Tele) Ue L 8

TRECHOPHYTON RAUBITSCHEKII SP, NOV.

JULIUS KANE
Medical Mycology Laboratory
Ontario Ministry of Health
Toronz.o, Ontario MSWeLR5S

TRA F. SALKIN
Division of Laboratories and Research
New York State Department of Health
Albany, New York 12201

IRENE WEITZMAN
Bureau of Laboratories
New York City Department of Health
New York, New York 10016

CATHERINE SMITKA
Medical Mycology Laboratory
Ontario Ministry of Health

Toronto, Ontario M5W 1R5

A urease-positive Trichophyton isolated from skin scra-
pings has been found to be sufficiently different from
existing taxa within this genus to warrant its establish-
ment as a new species, TIT. raubitschekii.

MATERIALS AND METHODS

TEST ORGANISMS - Forty-one isolates of a urease-
positive Trichephyton recovered from skin scrapings and
‘10 isolates Sane of T. (rubrum and 2. mentagrophytes
recently obtained from clinical specimens were used in
this study. The purity of all isolates was established
by single spore isolation and other methods (2,5,8). The
identification of T. rubrum and T. mentagrophytes
isolates was established by standard methods (6,9) before
they were selected for this investigation. All isolates
were maintained on peptone-dextrose agar (Sabouraud dex-
trose agar: 4% crude glucose, 1% Neopeptone, 2% agar)
at 26-28°C. Seven-day-old cultures on this medium served
as the source of inocula for all morphologic and physio-
logic tests.

MORPHOLOGY - The gross and microscopic morphology of
each isolate was observed after 2 and 4 weeks' incubation
on peptone-dextrose agar at 26-28°C.

GROWTH AND PIGMENT FORMATION - A pinpoint inoculum
from a 7-day-old culture was transferred to the center of

260

each agar slant. All slants were incubated at 26-28°C.
The extent of growth and pigmentation on bromcresol
purple-casein dextrose agar (5) and corn meal-dextrose
agar (3) were assessed 7 days after inoculation. The
extent of growth on lactose agar (4) was noted at weekly
intervals for 3 weeks.

UREA HYDROLYSIS TEST - Ability to hydrolyze urea
within 7 days was determined by the urea broth method of
Kane and Fisher (7).

HAIR PERFORATION TEST - Following the technique des-
cribed by Ajello and Georg (1), autoclaved human hair was
examined for wedge-shaped perforations at weekly inter-
vals for 4 weeks.

MATING REACTIONS - Mating tests with strains of
Arthroderma simii and A, benhamiae were performed accor-
ding to the method of Stockdale (10) on the oatmeal-salts
agar described by Weitzman and Silva-Hutner (11).

RESULTS AND DISCUSSION

Division: Fungi imperfecti
Form class: Hyphomycetes
Form order: Moniliales
Form family: Moniliaceae

Trichophyton raubitsehekii Kane, Salkin, Weitzman et
Smitka, sp. nov. Colonia in agaro peptone-dextrose,
postquam 14 dies apud 26°C lapsi sunt, 28 mm in diam.fit;
plana, tenuis, lenis usque ad granosum in textu, cum
centro fulvo sublatoque. Pars aversa erat sanguinea (#3
in charta colorum Raynor Mycology RMCC). Margo coloniae
erat distinctus cum angusta fascia sine colore.

Post 28 diebus, colonia erat 53 mm in diam.; plana,
tenuis, lenis, cum centro sublato exquo sulci radiantes
ad peripheriam pertinuerant. Tres zonae colorum notatae
sunt. Centrum erat submurinum, (RMCC #117) quod ad zonam
mediam, colore grisea-sepiaceam (RMCC #106) pertinuit
terminatumque ad aream peripheriam, lavendulo-griseam
CRMCC #125). Margo erat distinctus cum fascia sine
colore 3 mm in latitudinem. Pigmentum in parte aversa
Sanguineum (RMCC #3) remansit.

Sporulatio in agaro peptone-dextrose erat crebra et
in primis et in subsequentibus subculturis. Macroconidia
erant longa et angusta, 45-51 x 4,8-6.3 um, cum parieti-
bus teribus, fines obtusi, et ex 5-9 cellulis composita.
Microconidia erant in forma variabilia (clavata, sub-
sphaeralia, globosa), 4.8-6.4 x 3.2-4.8 um, aut sesselia
aut in brevibus conidiophoribus, a latere ferebantur.
Holotypus OMH 1094, teleomorph ignotum.

Habitat: Man
Holotype: OMH 1094, isolated from human skin scrapings,
Toronto, Ontario, Canada.

261

Living cultures of the holotype have been deposited
in the American Type Culture Collection, Rockville, Mary-
land (accession number ATCC 42631); the University of
Alberta Mold Hebarium, Edmonton, Alberta, Canada (access-
ion number UAMH 4314); and the Medical Mycology Labora-
Trorysculture, Collection, Ontario Ministry of ‘Health.
Orono, Ontaria, Canada (accession number OMH 1094). The
epithet T. rauhitschekii was chosen to honor the late Dr.
F. Raubitschek, dermatologist, medical mycologist, and
teacher.

The colony on peptone-dextrose agar after 2 weeks at
26°Cowas. 28emmiin diameten, flat, thin, velvety to
granular in texture, with an elevated, buff-colored
center (Figure 1A). The reverse was blood-red, #3 on the
Raynor mycology color chart (RMCC: Commonwealth Myco-
logical Institute, Kew, Surrey, England). The colony
margin was distinct, with a narrow colorless band.

After 4 weeks the colony was 53 mm in diameter, flat,
thin, velvety, with a raised center from which grooves
radiated to the edge (Figure 1B). Three color zones were
noted: a pale mouse-gray center (RMCC #117), a grayish
sepia (RMCC #106) intermediate zone, and a lavender~gray
(RMCC #125) peripheral area. The margin was distinct,
with a 3-mm colorless band. The reverse remained blood-
red (RMCC #3).

Sporulation on peptone dextrose agar was abundant in
both primary and subsequent subcultures. Macroconidia
were long and narrow, 46-51 x 4.8-6.3 um, with smooth,
parallel-sided walls, blunt ends, and five to nine cells
(Figure 2). Microconidia were variable in shape
(clavate, subspherical, or globose), 4¥.8-6.4 x 3.2-4.8
um, and borne laterally, either sessile or on short coni-
diophores (Figure 3).

Teleomorph unknown.

Pronounced urease activity was observed within 7
days of incubation at 26-28°C. Growth on bromcresol
purple-casein dextrose agar was restricted, and a reddish
pigment developed in the center of the colony. Brown
reverse pigmentation was observed on cornmeal dextrose
agar. Growth on lactose agar was restricted after 3
weeks' incubation.

, Mating of 12 isolates of T. raubitschekii with
strains of A. simii mating type (A) resulted in the for-
mation of either pseudogymnothecia or gymnothecia with
immature, irregular asci. In contrast, no reaction
occurred when the same isolates were paired with tester
strains of A. benhamiae or mating type (a) of A. simil.

The key morphologic and physiologic features of Ae
PubrUMS yh. raubacechekii, and 2. mentagrophytes are com-
pared in Table 1. Trichophyton raubitschekii is similar
to TI. mentagrophytes in urease activity but differs in
its inability to perforate hair in vitro, its restricted
growth on bromcresol purple- casein dextrose agar, and its
lack of sexual meaction with A. benhamiae. While resemb-
ling T. rubrum in its reddish “pigmentation on glucose-
peptone agar and sexual reaction with A. Simii (A),

Culture of i. raubitschekii on peptone-
dextrose agar at 26-28°C after (A) 2 weeks

(B)) Y weeks” incubation.

205

(A-H) on peptone-dextrose agar.

Macroconidia

Figure 2

Phase contrast, x 400.

264

T. raubitschekii differs from T. rubrum in four important
Gharacteristics: it is urease-positive; ite ferowthu.e
restricted rather than spreading on lactose agar; its
pigmentation on casein dextrose agar is brown rather than
red; and it produces abundant macroconidia.

Figure 3. Microconidia (A-F) on peptone-dextrose agar,
x 4OO0.

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266

ACKNOWLEDGMENT

We thank the regional Audiovisual Services of the

Ontario Ministry of Health for their assistance in the
preparation of the photographs; Ms. Margaret Kwok of the
Media Department, Laboratory Services Branch, Ontario
Ministry of Health, for preparation of experimental media;
Douglas Engstrom for help in the preparation of the Latin
diagnosis; and Dr. M.A. Gordon for his critical review
of the manuscript.

LEONA

alts

LITERATURE CITED

Avello, Li, and Lek Georges. 31957... sinavitromnaryr
cultures for differentiation between atypical iso-
lates of Trichophyton mentagrophytes and Trichophyton
rubrum. Mycopath.) Mycol. Apple 8:3=-1/.

Ajello, L., L.K. George, W. Kaplan, and L. Kaufman.
1966. Laboratory manual for medical mycology. U.S.
Government Printing Office, Washington, D.C.

Bocobo, F.C., and R.W. Benham. 1949. Pigment produc-
tion in the differentiation of Trichophyton mentagro-
phytes and Trichophyton rubrum. Mycologia) 41:291-
3022

Cowan, S.T. 1974. Cowan and Steel's manual for the
identification of medical bacteria, 2nd ed. Cambridge
University Press, London.

Fischer, J.B., and’ JU.*Kane, 1971. The: detection of
contamination in Trichophyton rubrum and Trichophyton
mentagrophytes. Mycopath. Mycol. Appl. 43:169-180.
Haleys#L.D. and C.S.-Callaway.) 1978. | Laboratory
methods in medical mycology, 4th ed. U.S. Government
Printing Office, Washington.e D.C.

Kane, J., and J.B. Fischer. 1971. The differentia-
tion of Trichophyton rubrum and *T. wwenigerepetes by
use of Christensen's urea broth. JwaMicrobiol.
Lyi O3.

Kane, J., and C.M. Smitka. A practical approach to
the isolation and identification of members of the
Trichophyton rubrum CVOUp-— LNs Proceedings vor avith
International Congress on the mycoses: superficial,
cutaneous, and subcutaneous. PAHO Scientific Publi-
cation; Washington, 9D.Cs 1980, (Sc sbubaeNno. 396
McGinnis, M.R. 1980. Laboratory handbook of medical
mycology. Academic Press, New York.

Stockdale, P.M. 1968. Sexual stimulation between
Arthroderma simii Stockd., Austwick and related
Species. Sabouraudia 6:176-181.

Weitzman, I., and M. Silva-Hutner. 1967.
Non-keratinous agar. media as substrates for the
ascigerous state in certain members of the gymno-
ascaceae pathogenic to man and animals. Sabouraudia
52335-3340.

MYCOTAXON

Val me Nett ee NOR me hee ma ty 1) Oy) 92.0727. Apri ledunes 9S.

CYRENELLA ELEGANS GEN. ET SP. NOV., A DIKARYOTIC ANAMORPH

S: E. GOCHENAUR

Department of Btology, Adelpht Untversity
Garden City, New York 11530

ABSTRACT

Cyrenella elegans Gochenaur is described and
phlustrated from sand around the base of the
mushroom, Laccarta truliteata (Bllis) Singer.

Lvs bright orange colony, bears) hyphaerwith clamp
CONNECTIONS , Small, obovate, dikaryotie conidia
with 3 or 4 apical appendages and a single basal
one. sandeclusters of elo boseschilanydoespores.
Monokaryotie blastospores may be produced. The
Osan SMe Ss .Orida Lay eames opi ewan dea cia dopiid ie;
Uo aZese Ni trate ws andmregud Nessie viLanins
p-aminobenzoate and thiamine.

INTRODUCTION

Several Rhodotorula-like isolates were obtained from
dilution plates prepared with sand scraped from the
submerged stipe of Laccarta trulltsata. One of these
colonies during subculture was found to produce hyphae with
clamp connections and small appendaged conidia. A new form
genus is introduced for this isolate as no fungus with
Similar characteristics appears to have been described in
the literature.

The holotype, a desiccated agar culture (No. 785), is
deposited in The New York Botanical Garden Herbarium (NYBG),
Bronx paN. 0) Yo. Living representative strains (isotypes) are
deposited in The American Type Culture Collection (ATCC),
Rockville, Md. and The National Culture Collection (NCC),
Commonwealth Mycological Institute, Kew, England.

MATERIALS AND METHODS

Monphological descriptions are based on a strain that
developed from a single conidium obtained from the orieinal
13O0labe.) “Lhissapkaryon and a cloned) blastosporic strain
derived from it were cultured on 2% malt extract agar with
2% glucose (MEA) at 20°C and examined microscopically
using brightfield and phase optics. Measurements of
MUCMOSeCOpreyoLrUuciumMes waremune mean Of ss Orderverminations (=
the standard deviation. COLlOrs rerermtvor pla less inetiie
Methuen Handbook of Colour (10).

Nutritional studies using the dikaryon and
blastosporic cultures employed a basal medium composed of
Bacto-Yeast Nitrogen Base without Amino Acids and Ammonium

268

Sulfate, (Difco Laboratonies, Chicago, 11.) andveoliditied
with Bacto-Purified Agar. This was+supplemen ted with
alucose (1G) sins tests ror wills ze tuonol vari Te aioe en
sources and with Bacto-Vitamin Free CaSamino Acids in tests
For UtalVzawion or, Verious .carben, Sources, All basal media
and stést, compounds. were Tilter. sterilized except compounds
LG throuen es CTable Iivowiveh were Neat cust lamenmat
121°C. Vitamin requirenents were determined torn cultures
grown on Bacto-Viteamin Free Yeast” Base solidified with
BBL-lon Agar No. 8 (Baltimore Biological Baboratori es,
Cockeysville, Md.) and supplemented with 0.2% Bacto-Vitamin
Free: Casamino Acids and the appropriate vitamins.
Vitamin-depleted inoculum was used.

Environmental studies, using the Dlastosporic) stage
only, were done in Jiqgudid culture employing a medium
composed of Bacto-Casaminos Acids (0557). anode slucose 172).
Growth was méasured spectrophotometrically at a wave length
of 450 nm. Additional temperature data were collected
using the, dikaryon erown) on MEA:

Chliamydospore germination Was studied using the
technique employed by Neweid. and) ela) (120" tor
Rhodosportdtum teleospores. Chlamydospores produced in
malt extract broth were collected, byicentritveationie waered
several times, resuspended in “sterile distilled water and
Svored Lor Up. to So months wate eoom Tempera Lim eee ind (OAS ©
Aliquots were removed at various times and streaked on
Bacto-VG Juice Agar and MEA, Simeubeted au 12°C and) 25°C.
and examined periodically over 4 days.

Nuclear number was determined using Furtado's
Toluidine Blue (5)... GCelblis were grown son MEA for) 2) to. 5
days 2b) 30°C and. then stained. iniwet mounvuss Ores -ar coe on®
eover glasses and pretreated, for +> Mine wito Lene lat. 60°C
before being stained.

TAXONOMY

CYRENELLA Gochenaur, forma-gen. nov., nomen anamorphosis,
Deuteromycotina, hyphomycetes pertains.

Mycelium ex. hy pDhiLs Ryalinice ramocic, =levyi ver
tunicatae compositum zygodesmatibus praeditae.
Contdtophona hyalana .macronemsa ta, solitvariac. vel
GCongregatae, erecta, irresulariter ramnosa. Cellulae
eOnsdiogpense monodlasticae, apiecalia jet) lateral iar
globosae, ovoideae vel cylindricae vel mixtae, vacuae’ post
dehiscentem conidiorum. Conidiea hyalina, obovate. oe cum
Drachiist apicealipus) Tredianti bus, 2-5 ou Ula ppend iver
Dasa iaei Chlamydosporae et blastosporae sint productae.

Species typica: Cyrenella elegans Gochenapr

ETYMOLOGY: Prom, the Greek, Cyurenes. a-mythicalwwa ter
sprite, an reference’ to the probable habitat or thes funeus
and. the-resemblance of ts) conidia to theses of #thesanuatiuc
hyphomycetes.

CYRENELLA ELEGANS Gochenaur, forma-sp. nov. Figs. 1-2.

Coloniae in agaro maltoso post dies decem ad 20°C
4-5mm diam. attingentes, humectae, calendulinae,

209

Sdinpactae, wn Cenurum acervulatum, margins planis, hyphis
aereis raris. Hyphae immersae leves, zygodesmatibus
praeditac, ad 2ilm crassae: Conidiophors hyalina,
macronemata, solitaria vel congregata, erecta,
mrregulariver ramoca, ad) 100uUm lonea, 3-4ium bateae. Cellulae
conldiogenae monoblasticae, apicalia et lateralia, ovoideae
mel cylindrvcae vel mixtac, vacuae post dehiscentem
conidiorum, 5-12um longae, 3-hum latae. UGH waahivee Isa nal,
mniceliularia,wornuecleolata., 10 x Vimy cum’ brachiis

S0tea lupus Pradiaget bus 3-4. 8L3-20m Joneis. Lum Latis ad
basen ct sO. >UmM la tis ad eap2cem, teu Uniappendici- eccentrica
Dac arike wy, =i. OO. Olims Chlamydosporae abundae, solitariae
et catenatae, intercalares et terminales, frequenter
gregarae, hyalinae vel melleae pallidulae, binucleolatae,
globosae vel subgiobosae leves et crassitunicatae,
5.5-6.5um Latae. pUuaLUuSs Siexualie Lenotis:

In arena, subs lacceaqrta trucilisatd,, Hempetead jake
bia ve: Park, Lone Leland. New fork, October, 1976s
HOLOtyDUSs “CULULUra (pura Siecata, (numero (05), NYBG Herb.

ETYMOLOGY: From the Latin elegans=graceful,
describing the appendages of the conidia.

DEVELOPMENT

Germination. of the conidiarot 0, felegans occurs within
24 hr on MEA at 20°C. One to 4 germ tubes arise apically
between the appendages and/or from the base, but rarely
from the lateral walls. Clamped septa and branches develop
where the germ tubes emerge from the conidium. As the
colony ages, conidia may germinate in place producing
apically, laterally or from the base, one or more
conidiogenous cells that develop conidia (Fig. 2C).
Cytoplasm flows into the conidiogenous cell leaving the
Original conidium empty. This process may continue through
several cycles, so that short chains of empty conidia are
common. AS a result of this proliferation, the color of
the central area of the colony is diluted and much lighter
than the periphery (Fig. 2A). Growth is slow. Colontes
attain a diameter of 3 mm in 7 da and approximately 24 mm
in 35 da. They are Persian to chrome orange (6A 7-8) with
a raised, rugulose central region consisting of pulvinate
masses of conidiophores and conidia and a narrow plane
Margin. With age, a broad, radiately wrinkled periphery
composed of mycelium and chlamydospores develops (Fig. 2A).
It is somewhat fiberous in appearance with small mounds of
sporulating cells scattered over the surface or coalesced
to form larger irregular or wedge-shaped patches. The
surface is moist. Aerial hyphae are rare or absent; when
present they are usually aggregated into tufts. The
myceltum is hyaline and consists of sinuous and straight
hyphae of uniform width, 2.0+0.3um diam, with smooth,
Slightly thickened walls. Septa with clamp connections
dome b eb Lrot OCCUL approximately Joa llim Prom theacip
and then’ at_intervals of approximately 507m. The cells are

binucleate. A dolipore apparatus was not revealed in
preparations stained with ammoniacal Congo-Red (11).

21-0

Us ieet ee Cyrene lia, elegans’.
Cr Con dilact (RB) Stages wp tinetgcve lopment aor

GonidiLogenous celts.

and clustered chlamydospores.

of cytoplasm.

RACAL

Adventitious simple septa are common in older hyphae and
coralloid or haustorial branches similar to those shown by
Otaeve (3) forvlt lobastdzum may occur.

Contdtophore development begins within 2-3 da. The
conidiophores arise as erect, branched hyphae
indistinguishable at first from the vegetative cells. They
are hyaline, of variable length and produce thumb-shaped
terminal and lateral branches that stain intensely with
basic dyes (Fig. 1B). These branches are separated from
the conidiophore early in development by clamp connections.
They also may be produced individually along the vegetative
hyphae (Fig. 2B) or less commonly from a clamp. They may
differentiate directly into conidiogenous cells or may
branch once or twice, each branch becoming a conidiogenous
cell separated from its parent cell by a clamped septum.
The irregularly branched conidiophore at maturity (Fig. 1C)
is generally less than 100um tall, 3-4um wide at the base
and is best observed using phase contrast optics since its
upper half consists of cells devoid of cytoplasm and
somewhat collapsed. Conidiophores are often aggregated
into sporodochial-like clusters that arise from a basal
mass of intertwined hyphae and chlamydospores. The
conidiophores produce single or clusters of 2-3
conidiogenous cells laterally and terminally along their
branches. The contdtogenous cells are phialide-like,
binucleate, have a thick-walled body and a thinner-walled
neck that is approximately 1-4um long and 1-2um wide. They
vary in form from globose to ampulliform to cylindrical and
measure 2-4um wide, 5-12um long including the neck. Each
produces a single conidium.

Mature conida appear within 4-5 da. Contdta (Fig. 1A)
are blastic, orange in mass but hyaline by transmitted
light, binucleate (Fig. 2C), obovate, thin-walled, 10 x 4um
(eit. 2 x 4.24 5m) and bear 3-4 long,, narrow, divergent,
flexible, apical appendages, 16+2.4um long, lum wide at the
base and tapering to 0.5um wide at the apex, and an
Sccentric paca lmappendagersor suntftormewiath, 96 9F2 alex. Ooum.
At 20°C, approximately 33% of the conidia examined had 3
apical appendages; 66% had 4. At temperatures above 25°C,
aDOULeLSeeOf the conidiashad one, 27 and)5@arms:.

Chlamydospores (Figs. 1D, 2D) appear after 3 da. They
are Single and catenulate and arise terminally, intercalary,
or less commonly from clamps. They typically form in large
compact clusters behind the margin of the colony and just
above or below the agar surface. They are subglobose to
globose, 6+0.4um diameter, binucleate, hyaline to pale
vamber, thick-walled and filled with guttulae. Single ones
are often oval and/or unsymmetrical and borne between
several adventitious septa that form as the cytoplasm
retracts into the developing chlamydospore.

Chlamydospores germinate within 3 da at 20°C (Fig. 2E).
A single germ tube that branches and develops clamps where
it exits from the spore is produced and a typical colony
with conidia develops within 5 da. Chlamydospores remain
viable after storage for up to 5 mo at room temperature in
distilled water. Those held at 7°C exhibited a progressive

FU. 2 Cyrenella elegans.

(Ny) Colony on melt extract agar after 35 da. at ish deus
(B) Vegetative hypha with a clamp connection. Arrow

designates a young conidiogenous cell. (C) Binucleate

Arrow designates a young conidiogenous cell and

conidia.
situ germination.

secondary conidium produced during. in
Three empty conidia appear on the Left.

decrease in the percent germinating after 45 da storage and
showed no germination after 5 mos. Karyogamy, probasidia
and soredia were not induced by these conditions, nor do
these spores resemble, in either coloration or mode of
formation, the teleospores produced by members of the
genus Rhodosportdium (2, 3, and comparison in culture with
Peemcacryovaum Kells, Hunters, Tallman ATCC 24502. & ATCC
24503). In view of the above, it is considered likely that
these structures are true chlamydospores and not the
teleomorphic state of ¢. elegane.

Dedikaryotization occurs spontaneously, but
infrequently, as the culture ages; a uninucleate yeast
phase results. It can be detected readily by streaking
material onto an agar medium containing raffinose, a sugar
utilized less well by the filamentous stage. Origin of the
blastospores is unknown. Subjecting the dikaryon to
flooding with water or anaerobic or microaerobic conditions
while growing it on media with various C/N ratios and low
or abundant in nutrients failed to consistently induce the
yeast phase.

The yeast colontes, like the dikaryon, grow slowly,
reaching a diameter of 8-10 mm on MEA at 20°C after 30 da.
They are slightly elevated when viewed in cross section and
show faint radial lines. Their surface is semi-glossy,
deep orange to Persian orange (6A8-7A8) in color with a
Slightly deeper carrot red reverse (7B7). The yeast cells
(Fig. 2F-G) are unicellular, monokaryotic, non-encapsulated,
elliptical to cylindrical with rounded ends, less commonly
epclava leo .0+) , 86x, 3..2+0. 81m, andsreproduce, by terminal
and subterminal budding. Fermentation is absent. In ME
broth, most cells are single and only rarely are short
chains of cells observed. No pellicle is produced.

Evidence suggests that the nuclei in the dikaryon are
identical. A dikaryophase is not reestablished when yeast
cells from randomly selected colonies are mixed on cornmeal
agar. However, it develops spontaneously in every cloned
yeast colony that is over a few days old. Cells in the
center of a colony at the agar surface become binucleate.
It is uncertain whether these occur following conjugation
between two cells or by failure to bud during mitosis.
Observation of empty single yeast cells with an attached
binucleate conidium suggests that the latter is the case.

A sparse mycelium always develops under every yeast colony
but its further expansion is limited unless the overlaying
cells are scraped away. All cells on the surface of a
colony remain monokaryotic so that the yeast phase can be
maintained indefinitely in culture but always contaminated
eventually with the dikaryon.

(D) Binucleate chlamydospores. (E) Germinating
chlamydospore. (F) Budding blastospores. (G) Uninucleate
blastospores. Material in photographs B-G was stained with

Coluidine, blue. Cells in photograph G were pretreated with
Pench tore Se minemastu60 2C,

274

NUTRITION

C. elegans is an acidophilic, mesophilic, obligate
aerobe. Growth occurs over a wide range of H-ion
concentrations. It is best between pH 3.5-6.0 and absent
below pH 2.5 and above pH 7.4. Conidia do not germinate at
5°C and 37°C. Pigmentation iS more intense and colonial
and cellular development best between 18-23°C. Growth is
fastest at 30°C; absent above 34°C.

Table I. Growth of Cyrxenella elegans on various nitrogen
sources®

Compound Growth Compound Growth
1., Potassium, nitrate + 6.9 Glutamic acid +
Cot POLAS SLUM Ww iat ra te ~ [. Asparagine +
3. Ammonium sulfate + G7 ) Casamino tacids ++
4, - Urea + O*e G eutart in. ++
Sp araspearct cea Cand as 20. ‘Sodium Caseinate ++
a)

Cultures: were Aancubated fore5 da. ate co. Cc.

+t Growth equal to growth ‘on caseamino acids

+ Growth weak (but, better “thanyecrowth on’ a mediumplacking
a nitrogen source

- No growth

C. elegans utilized all nitrogen sources tested with
the exception of potassium nitrite (Table I). Growth and
development are best on a mixture of amino acids. The
proteins, gelatin and sodium caseinate, are utilized but
diffusible hydrolases were not detected when colonies on
these substrates were flooded with acidified HgCl.,. A
variety of compounds can serve as the carbon-energy source
(Table II). Growth and development are best on glucose or
fructose. Krebs Cycle acids (Nos. 37-38) are assimilated
but not compounds with ester linkages (Nos. 23-24). Among
the polysaccharides (Nos. 17-22), only the fructosan
inulin and the starches (Nos. 18-19) were weakly degraded.
Diffusible exoenzymes were not detected when colonies
growing on the latter compounds were flooded with an IKI
solution. Patterns of carbon and nitrogen assimilation by
the blastosporic and dikaryotic stages are identical. The
exogenous vitamins, p-aminobenzoate (PABA) and thiamine,
are required for growth of both the mycelial and yeast
phases. High concentrations of folic acid can be
substituted for PABA but not a mixture of amino acids and
purines.

DISCUSSION

Cyrenella elegans is a member of the Basidiomycotina
because it produces mycelium with clamp connections but the
absence of a telemorphic state makes its phylogenetic
position in the subdivision uncertain and any attempt at
ordinal placement a purely speculative venture.

Peple Il. .Growth of Cynzenella pedal on various
carbon-energy sources®

Compound Growth Compound Growth
ie. Ay Lose plat ian.) ae Gh ~
Oe. L-Arabinose ak ane DNA -
3. Ribose - 24, Triglyceride -
Wa Glucose ++ 25. Fucose =
5. Fructose ++ 26... L-Rhamnosce ++
6. Mannose ++ 27 Glucosamine =
Galactose - Oo ein thane | -
8. L-Sorbose ~ 29. Glycerol ++
9. Trehalose + 5 Oren Gye ILE? 166.0 -

A) Sucrose aap Biles DOr Ost OL ++
Thane Maltose tacts Bier Mannitol Par
ieee Melibiose - Se Lnositol -
ieee Cellobiose + Si eeeGalactitol -
mo i Lactose a 35. Acetate =
15. Melezitose ++ 26... DL-Lactate =
fore Raftinoce + 237. Succinate ++
igs snl + 3.0 eC Lo GaLe +
18. Amylose + 30 ee Dar Gen “=
19. Amylopectin + WOe 4oal Loan +
20. Pectin - i Vandel n -
mits Celiulose z. re. Casamino acids =
ayaa compounds were D-isomers except where noted. Cultures

onwcompounds 17-24 were incubated for ll «da at 126°C van.
Gunersmcor)> sda at oo 7 ek

tin.rOow Oredila lato Om bevteratian ~2rowth "on —slucose

+ Growth weak but better than growth on a medium lacking
a-carbon source

ea bOwtd meCduad le tLOuOr Less (than growth on .a medium Lacking
a carbon .source

The blastosporic stage could be readily accommodated in
the genus Rhodotorula (14) if the dikaryophase were lost.
It resembles R. aurantiaca (Saito) Lodder and R#. lactosa
Hasegawa in producing cylindrical cells, in colony color,
in utilizing nitrate, cellobiose and maltose and in
requiring PABA and thiamine. It differs from the former in
being galactose negative and L-rhamnose and raffinose
positive and from the latter in being unable to utilize
melibiose and lactose. The requirement for PABA is confined
to a very few wild-type fungi, certain species of
Rhodotorula, Blastoecladta pringshetmit Reinsch and single
strains of Saccharomyces cerevtstae Hansen (4). Because
this requirement is so uncommon, there appears to be strong
selection pressure against such fungi in nature. Its
Simultaneous occurrence in Rhodotorula-and Cyrenella may
Signify a much closer relationship between these genera
then is evident simply on the basis of similarities in
morphology and nutritional patterns.

C. elegans was obtained from sand along the shore of

276

a fresh water lake in an area that is regularly submerged
each spring for several months. Even though it came from

a terrestrial habitat, C. elegans has several
characteristics in common with organisms isolated from
aquatic environments. Its thin-walled, tetraradiate-like
conidia bear appendages in a pattern reminiscent of that
evolved by many fresh and a few salt-water fungi. They are
remarkably similar to the basidiospores of the marine
fungus Wia in size and shape but not development (9) and
superficially resemble the conidia of aquatic hyphomycetes
belonging to the genera Acaulopage Drechsler, Clavatospora
S. Nilsson and Clavartana Nawawi. Diffusible exoenzymes
are a characteristic feature of terrestrial fungi (1, 8)
but would be a distinct disadvantage for an aquatic
organism since such proteins and their digestion products
would be lost into the water. C. elegans produces no
diffusible hydrolases when utilizing starch, gelatin and
casein. Since its hydrolases must be retained in
protective association with the cell wall, it would be well
adapted for life in dilute aqueous environments. Finally,
C. elegans' requirement for PABA occurs less frequently
among terrestrial fungi than among aquatic ones. In

R. auranttaca, for example, only strains of marine origin
demonstrate an absolute need for PABA (14). These features
suggest that C. elegans is not a terrestrial fungus but is
an aquatic or possibly an amphibious one, whose propagules
were trapped in the sand as the lake water receded,
persisting there until isolated several months later.

If Cyrenella is an aquatic genus, it is an extremely
uncommon One., Dr. R.«Bandonivof the Univers1rty.of British
Columbia (personal communication) reports that C. elegans
appears to be identical to a fungus he collected from a
driftwood chip on the shore of the Iowa River about 25
years ago. Other workers who have examined the conidia
report they do not remember seeing spores of exactly this
type in foam samples nor have illustrations of these spores
appeared in the many works describing propagules from this
environment (6, 7). With its slow growth rate in culture,
superficial resemblance to Rhodotorula and tiny conidia,
it is understandable why a Cyrenella could be overlooked.

ACKNOWLEDGMENTS

LaWisen to tTheank tHestollowings individuals For their
aids Dre sR. Bandoni forsstudying cultures, or CUsenc Cede ond.
offering many shelpPrul suggestions as welbtlas kindly
YPEVIieWwing the umanuscripts Drs. W. Bridge Gooke, J. Leland
Crame, C. Tt -dngold, Rs H. Petersen; and yshun-—1ehi Udacawa
hor Pe ranmining slides eop thestuneue:.0r. 4c. el . hogereconet or
assistance with they taxonomy; Dr. A. H. Bréenowitz ior
takpnesthe photomicrographs: Prot. Lh. Ascher, language
Dept. Tonecorrectine the Latam diagnosis: Ms .Julie
Anderson, a former undergraduate in my laboratory who
studied Cyrzenella with me as part of her honors thesis; and
Mrs. W. Jaworski for typing the camera-ready copy.

Dhal

LITERATURE CITED

meee Domsch, K., and Ws Gams. 1969. Variability -and
Pocemtial of a souid fungus: population to decompose pectin,

xylan and carboxymethyl cellulose. SOid Biol. Biochem.
me (29-36.

Pree eed a ce Phath and S.aNewell. BS yy
Rhodosporidium Banno. pp. 803-814. In: The Yeasts. Ed.,
ae LOUder. end ed. North-Holland Publishing Co., Amsterdam.

Cree ce Ll Sais else pHUR Ger, and. A. Valiman’. eae, ar Pie
basidiomycetous.yeasts (Rhodosporidium spp.n.) with
setrapolar and multiple allelic bipolar mating systems.
Pied | Microbiol) )19: 9643-657.

Lig Fries, N. 1965. The chemical environment for fungal
growth. 3. Vitamins and other organic growth-ftactors.
Pee t91-523- In: The Fungi. _Eds., G.” Ainsworth ‘and
MmEicocman. Vol. I. Academic Press, N. Y.. N.Y:

i. ia Sh Bee Me Me ees Pa iar 1OTO.. Atconol@ te LOolvidine vplue: A rapid
Memo LOr staining nuclei an upiixed mycevozoa and fungi .
Mycologia 62: 406-07.

6 Mascot ds Cx. NEO TG Advances in the study of so-called
aquatic hyphomycetes. Amer. J. Botany 66: 218-226.

foeericOld, ©. bO1>. ~ Guide to aquatic hyphonycetes:
Freshwater Biol. Assn., Ambleside, England.

‘eae Cyol Per A, and=p .) -aStruwe. LO OF. Microtung i 16 f
decomposing red alder leaves and their substrate
Peabication.) SoriyBiol. Biochem. 12:7" 425-431.

OF Kohimeyer, J., and E. Kohlmeyer. LOO. Marine

mycology the higher fungi. feademiouPpress . Nia ey. seth ede
690 p.

Move kornerup, A.;, and J. Wanscher. POOTE Methuen
pemovook Of colour. 2nd ed.) Methuen and Cos, )Ltd., london.
Uns = p's

Jil. Nawawi, A., J. Webster and R. Davey. 1977.
Dendrosporomycees prokifer gen. et sp. nov., a basidiomycete
Birhsbranched conidia. Trans. Brit. Mycol.’ Soc. 68: 59-63.

mer Newell. S., and J. Fell. LO FOR The perfect form of a
marine-occurring yeast of the genus Rhodotorulka
Myeotogia 62: 272-261.

Ss Oa Wei, 5 ste 1968. An unusual new heterobasidiomycete
with Tilletia-like basidia. pp. 261-266. In: Mycological
moumtes honoring John N. Couch. Ed... We J: Kock. The Univ.
Peecarolina Press, Chapel Hill.

Beeeenaff, H., and D. Ahearn. 1971. Rhodotorzufla Harrison.
Peeelilof-1223) “In: The Yeasts. |Ed. J. Lodder. 2nd ed.
North-Holland Pub. Co., Amsterdan.

MYCOTAXON

Vol rex Vile N Ota. ap Demet OF 776.0 Apr un eeeho & 1

REVUE DES LIVRES

par

G.L. HENNEBERT
Book Review Edttor, Crotx du Sud 3, B-13848 Louvatn-La-Neuve
Belgtque

GENERA OF HYPHOMYCETES, par J.W. CARMICHAEL, W.B. KENDRICK, I.L.
CONNERS and Lynne SIGLER, 390 p., 129 pl., 8°, reliure spirale,
1980, The University of Alberta Press, Edmonton, Alberta, Canada.
iD aalp ae jeuc all Frosh

Cet ouvrage est une seconde édition entiérement revue et complétée
de la premiére psésentation par les premiers auteurs d'une compilation
illustrée et critique de la littérature originale des genres de Hyphomy-
cétes dans The Fungt de Ainsworth, Sparrow and Sussman, Tome IV A, en
1973. Les auteurs ont recensé cette fois plus de 2000 noms de genre,
contre 1541 en 1973. Les 61 planches de figures d'alors sont devenues
129 aujourd'hui. Aprés une introduction sur la méthode et la classifi-
cation des types conidiogénétiques, on y trouvera une liste documen-
tée des noms de genres, avec date, référence, type morphologique, hdte,
espéce type, statut nomenclatural et synonymie, avec références. Cette
liste distingue les genres acceptés par les auteurs des noms rejetés ou
douteux. Viennent ensuite les illustrations, soit reproduites des des-
criptions originales, soit dessinées a partir de publications récentes
ou encore du type. Les 853 figures représentent pour la plupart les
espéces types des genres et sont identifiées a l'espéce. Le grossisse-
ment n'est pas donné mais n'est pas indispensable a ce niveau. Suivent
encore 5 index: un index des genres par type conidiogénétique, un index
des connections entre téléomorphes (stades sexués) et anamorphes (stades
conidiens), une liste bibliographique, une liste des abréviations et
une liste des figures. Les auteurs ont suivi la nomenclature illégale
basée sur Persoon, Synopsts Fungorum, de 1801 comme point de départ,
contrairement a l'art. 13, mais combien souhaitée par beaucoup. Le livre
remplace les guides des genres publiés jusqu'a présent. Il est fort bien
imprimé. Sa reliure en spirale s'inspire de celle du Barnett, Illustrated
genera of Imperfect Fungt, mais on pourra le regretter.Vu son évidente
utilité et son prix assez modique, le livre aura sans doute une large
diffusion.

Les auteurs l'ont dédié a la mémoire de Luella K. Weresub, mycolo-
gue du Biosystematics Research Institute d'Ottawa, tant appréciée de
OUGT

CHAMPIGNONS D'EUROPE, par M. SVRCEK et J. KUBICKA, adapté en fran-
Gais avec coll. de G.L. HENNEBERT et al...296 p., 448 ph. col-,

8°, cartonné, 1980. Ed. Elsevier-Sequoia, B-1940 St Stevens-Woluwe,
Belgique.

Ce guide des champignons est remarquable par le grand nombre d'
espéces décrites - 448 - chacune convenablement illustrée d'une photo
en couleurs dans le milieu naturel. Les descriptions sont précises

209

et donnent les dimensions sporales. Une introduction illustrée retrace
la structure des champignons et leur classification, donne les conseils
au récolteur et le documente abondamment sur la toxicité des champignons.
Offrant un éventail exceptionnellement large des champignons d'Europe,
tant de montagne que de plaine, ce guide surpasse un grand nombre
d'autres moins complets.

ECOLOGY AND DISTRIBUTION OF FUNGI, Scripta Mycologica n° 9, 144 p.,
12°, broché, 1980, Academie des Sciences d'’Estonie S.S.R., Tartu
MstonLe.errix., RD il, LO.

Ce fascicule est intitulé en fonction des deux premiers articles
quil comprend : The composttton and seasonal dynamtes of the fungal
cover on mtnera sotls and Trophte groups of Estontan Agartcs, par K.
Kalamees. Dans le premier travail, l'auteur reléve les flores d'agari-
cales dans divers types de foréts et de prairies jusqu'a la tourbiére
et le marais et note sa composition et sa richesse spécifique 4a chaque
écotype végétal. Dans un second article, il regroupe les Agaricales
en 13 groupes trophiques, les mycorrhizogénes, les saprophytes de divers
types de substrat et les parasites étant les principaux d'entre eux.
Le fascicule se termine par une revision du genre Lastobelontum par A.
Raitviir, ot 1l'auteur combine diverses espéces et en décrit cing nou-
velles. Texte en anglais.

THE CHEMOSYSTEMATICS OF THE LICHEN GENUS PERTUSARIA IN NORTH AME-
RICA NORTH OF MEXICO. par Martyn J. DIBBEN, Milwaukee Public Mu-
seum, Publications in Biology and Geology N). 5, iv + 162 p., 136
fig., 4°, broché, 1980. Milwaukee Public Museum, Milwaukee, Wi.
DoZo Sun UOA Prix fy 220 te postage.

L'examen de pas moins de 6800 specimens, incluant entre autres
des matériaux historiques de quelques 40 herbiers reconnus, a été né-
cessaire 4a cette revue des 66 espéces North Américaines des lichens
du genre Pertusarta. 14 espéces sont nouvelles. 33 espéces sont endémi-
ques a 1'Amérique du Nord. Du point de vue de leur distribution écologi-
que, 60 2 spnt tempérées, 35 % arctoboréales et 5 4 tropicales. 40 acides
organiques sont recensés chez les Pertusarta, L'auteur a appliqué 1'
analyse chromatographie sur silicagel a tous les échantillons, selon la
méthode de Culberson. Sa taxonomie des espéces est avant tout basée sur
la morphologie du thalle et de la fructification apothéciale. Sur cette
base elle distingue deux sous-genres, P, sbg. Pertusaria et P.sbg.
Ptonospora, l'un a fructification amplicariale ou pertusariale, l'autre
a fructification lécanorale ou sorédiale, L'analyse chimique s'avére
une confirmation précieuse dans la distinction des espéces. Confrontées
entre elles, les deux méthodes permettent de réduire le poids donné aux
variations et de ne plus subdiviser l'espéce en une série de variétés
ou de formes, La différenciation chimique jointe 4a la ségrégation géo-
graphique a permis cependant de confirmer des espéces proches comme dis-
tinctes. L'espéce type Pertusarta pertusa, eurasiatique, n'est pas recon-
nue en Amérique du Nord, celle qui lui a été confondue est décrite comme
P. consoctans. L'auteur a le souci d‘une nomenclature correcte. Elle
indique le nom du genre comme nomen conservandum, notant qu'il est illé-
gitime (Art, 63), et référe sans plus de détail - A regret - 4 un article
futur. Ses synonymies précisent lestatut nomenclatural et sont justifiées
par des commentaires, Les descriptions des espéces d'une liste détaillée

des spécimens examinés et d'une distribution géographique illustrée. La

280

technique macrophotographique appliquée laisse a désirer. Cette monogra-
phie 4 la fois nouvelle, précise et didactique mérite 1!'attention:

A PRELIMINARY POLYPORE FLORA OF EAST AFRICA, par Leif RYVARDEN &
Inger JOHANSEN, 64° °p., 212 fig. , 8°) couverture papier, 1980,
Funghtlora: Ed’ -P50.Box 95") Blindern, Oslou3, orway.s (PP) ix, Nka 200;

Durant les 10 ans passés, alors qu'il publiaita intervale régulier
les volumes des flores des Polyporaceae et des Corticiaceae d'Europe du
Nord (revus dans Mycotaxon), le premier auteur a élaboré cette flore
des Polypores' d'Afrique orientale par diverses explorations de récol-
te et 1'étude des grands herbiers européens. La flore couvre 5 familles,
les Corticiaceae, Ganodermataceae, Hymenochaetaceae, Polyporaceae et
Tremellaceae, 75 genres dont 4 nouveaux (Antrodtella, Echtnoporta, Navi-
sporus et Pseudoptptoporus) et 337 espéces dont 3 nouvelles. Des clés
réellement dichotomiques des genres et espéces sont poposées et, évidem-
ment (et pourquoi?) limitées aux seuls genres et espéces traitées. Les
espéces sont décrites avec une attention particuliére aux détails anato-
miques et microscopiques, lesquels sont dessinés dans la plupart des
espéces. L'habitat et la distribution géographique sont indiquées, mais
sans liste des spécimens étudiés (le livre se veut @6tre une flore et non
une monographie). Imprimé en caractéres dactylographiques, le texte est
sans relief, peu attirant. Les nombreux espaces blancs donnent au livre
une épaisseur de 35 mm qui fera rapidement céder la faible couverture
de papier. Le livre est dédié 4 la mémoire de M.A. DONK.

SAPROPHYTIC MICROFUNGI FROM TAIWAN. PART 1 HYPHOMYCETES, by Takashi
MATSUSHIMA, Matsushima Mycological Memoirs No 1, 82 p., 46 fig.,
8°, paperback, May 1980. Publication Matsushima Fungus Collection,
23-19-6001 Mikageyamate-2-Chome, Higashinada-ku, Kobe, Japan 658.
Price: free of charge trough official request.

This is the first of a series of papers enumerating the microfungi
collected in Taiwan during 1976 to 1978, by the author when he was guest
as visiting senior mycologist at the Plant Pathology Division, Plant
Protection Center, Taiwan. As in previous publications, the authors
reports his profuse collections and cultures of the Hyphomycetes with
the same wonderful ability and efficiency. His descriptions or comments
on species are just what is needed, but his ful page drawings speak more
than words. Not less than 250 Hyphomycetes including 2] new species and
2 new genera(Acumtspora and Chetropolyschema) are recorded, documented
and 46tillustrated on the 82 pages of this issue. This is the start of
a new mycological series entitled Matsushtma Mycologtcal Memoirs. We
are sure of the succes which this series will receive through the
mycological world. This series is distributed by the author's newly
established institution, a very well standing fungus herbarium and cul-
ture collection, the Matsushtma Fungus Collectton.

BIBLIOGRAPHIA BOTANICA CECHOSLOVACA 1973-1974, par A. Neuhaduslova-
Novotna et D. Guthova-Jarkovska, 564 p., broché, 14x20 cm,Botanicky
Ustav GSAV, Prithonice u Prahy, 1978.

Ce répertoire bibliographique donne toute la littérature botanique
publiée en Tchéchoslovaquie durant la période citée, classée par matiére
(Sle ieee EHONEEUNE

———e

281

UNTERSUCHUNGEN ZUR KONSERVIERUNG DE FRUCHTKORPER DES SPEISEPILZES
PLEUROTUS OSTREATUS UND DER PARTIELLEN AUTOLYSE VON PILZSELLWANDEN,
par Helga SCHMITZ, Bibliotheca Mycologica vol. 77, 98 p. Se yidos
papier, 1980, J. Cramer ed.

Dans 1'étude du mécanisme enzymatique de 1l'attendrissement observé
lors de la conservation de carpophores dans des solutions acidulées
ac€tique, citrique et lactique (pH 4) l'auteur met en évidence 1'action
de R-glucanase et chitinase spécifiqueSdu pied et du chapeau du fruit,
tandis que les S-glucanase et chitobiase du mycelium sont sans action.
Thése réalisée sous la direction du Pr. G. Eger-Hummel, Marburg.

MUSHROOMS OF IDAHO AND THE PACIFIC NORTHWEST. DISCOMYCETES, par
Edmundtine TYLUTRIs 133° pe9027) £10.45 80 phot. =) Loxzlem, 1979;
University Press of Idaho, University Station Box 3367, Moscow,
Idaho 83843, USA.

Ce livre est écrit pour les amateurs "enthousiastes" qui veulent
dépasser les fontiéres des basidiomycétes et s'aventurer a récolter les
morilles, helvelles, truffes et autres grandes Pezizales, Tubérales et
Geoglossacées. Chaque espéce décrite est illustrée d'une photo blanc-
noir. Une clé "de terrain, purement macroscopique" est proposée: mais
elle est dichotomique et couvre 5 pages bien pleines... qu'il faut lire
a chaque récolte? Une clé synoptique serait plus didactique et plus
rapide. Cette remarque dépasse le cadre de ce livre qui est bien écrit
et agréable. Pas moins de 122 espéces y sont classées et reconnaissables
par les clés,

CBS COURSE OF MYCOLOGY, par W. GAMS, H.A. VEN DER AA, A.J. VANDER
PLAATS-NITERINK, R.A.SAMSON, J.A. STALPERS, 2e édition, 110 pe,
JOR tL OM einaG sedosepaprer, —ChS\, baarn, Nederland. 19802

Le texte de ce cours (le édition revue dans Mycotaxon 3:558, 1976)
est revu et complété de la littérature récente, sous a peu prés le méme
volume. Les auteurs y développent les traits généraux de la morphologie
des champignons, dans chaque groupe taxonomique, étayé de nombreuses
figures et références. Le fascicule ne se veut pas un guide d'identifica
tion (voir von Arx, Champignons sporulant en culture pure, 3e éd. en
prép.) mais une bonne introduction au régne des champignons.

A REVISION OF CHRYSOSPORIUM AND ALLIED GENERA, par C.A.N VAN OORS-
CHOTeGStudiesiin Mycology sin -@20,289Fp. 86 fiptes oi ple.eines sados
papier, 1980, "CBS. eBaarn, Nederland.) Prixghti25.-%

Cette monographie donne les descriptions illustrées et les clés de
Chrysosportum (22 espéces), Myceltophthora (8 espéces), Emmonsta (2 var.)
Zymonema (1 esp.) Trtchosportella (2 esp.) et Geomyces (3 var.). La coni-
diogénése blastique et thallique de ces champignons est décrite, ainsi
que les relationsavec les stades téléomorphiques appartenant aux Gymno-
ascaceae, Onygenaceae, Ascosphaeraceae et Sordariaceae. Cette revision
est la premiére depuis le travail de Charmichael de 1962.

EESTI SEENTE KOONDNIMESTIK ( LIST OF ESTONIAN FUNGI, WITH HOST INDEX
AND BIBLIOGRAPHY) par J. JARVA et E. PARMASTO, 331 p., Din, dos
papier, 1980, Institute of Zoology and Botany, Academy of Sciences
of thesestonianis. so. Rolantu. pEstouiaraulrixcear Dl vee sha

Boe

Cette liste des champignons de 1'Estonie est le résultat de 15 ans
de travail des mycologues de 1'Académie de Tartu pour le dépouillement
de 1200 publications citées en annexe. Chaque espéce est citées avec hé-

tes et références.

BIOLOGY IN THR BLUE RIDGE, Fifty Years of the Highlands Biological
Station, 1927-1977, by Ral piisMs,SARGENL. #l>6up ae Liao a epapeien
back, 1977, Highlanas Biological Station, P.O.Drawer 850, High-
lands N.C. 28741.

The then named Highlands Museum and Biological Laboratory, initia-
ted in 1927 by a few determined people eager to preserve local human and
natural history»has been, from the early years, associated with mycology.
The Highlands station, in 1934, received the Mycological Society of
America for its summer foray. Dr. William Chambers Coker, who presided
the institution from 1933 to 1943,and subsequently as Honorary President,
has been, with Dr L-R. Hesler, Vice-President, the promotors (of ‘the’ stu-
dy of the Appalachian fungi. Coker published on basidiomycetes, parti-
cularly Hydnum and the Boletaceae (1939-1951). Hesler continued his Wotes
on Southern Appalachtan Fungt in Mycologia from 1936 to 1955, and colla-
borated with Alexander H. Smith on Hygrophorus and Lactarius (1932-1962).
Dr. L.S. Olive (1943-1953) published on Tremellales and in 1965 his book
Potsonous, Edtble and Halluctnogente Mushrooms. Dr. Ronald H. Petersen,
more recently investigated the clavaroid fungi and the aquatic hypho-
mycetes (1962-1963). Surely, the Blue Ridge Laboratory contributed much
in 50 years to the American mycology. The Sargent's book, retracing,
years after years, the multiple ways and reviving the unique spirit of
the Highlands Biological Station will please the present and the future
scientific people.

PHYTOPHTHORA CINNAMOMI AND THE DISEASE IT CAUSES, by G. A. ZENT-
MYER, Monograph No 10,-96 p., 39) figs’,°4°, paperpack,1980, The
American Phytopathological Society, 3340 Pilot Knob Road, ‘St’ Paul
MNS D2 errr cetUses: Oar

How is it possible to cover such widespread and largely investi-
gated fungus pathogen like Phytophthora cinnamomt in 95 pages. The author
did, and on the base of 600 literature references. The pathogen is of
importance for the decay of many tropical and temperate crops and trees.
900 hosts have been listed. The monograph details the many aspects of
the biology, ecology, physiology and genetics of the pathogen, all data
necessary in the search to. an effective control. The text is dense
and illustrated of many original documents. George A. Zentmyer of the
University of California, Department of Plant Pathology is devoting
himself on the study of that pathogen for already 20 years and published
with his collaborators more than 100 papers onthe subject. His master-
ship in that fidd will help many people engaged in research through the
world.

A MANUAL OF ASSESSMENT KEYS FOR PLANT DISEASE, by Clive JAMES,
44 fiches illustrées, 10x18 cm., 1980, The American Phytopathologi-
ealisociety 7, *StePatl< eprixrUsens 1020.

Ces fiches illustrent les clés d'évaluation des dégats par Rhyncho-
sportum, Puecetnta, Septorta, Drechslera sur céréales,Phoma, Leptotrochila
Stemphyltum sur luzerne et tréfle, Phytophthora, Streptomyces, Rhtzocto-

283

nta sur pomme de terre et Xanthomonas sur haricot. Elles aideront les
phytopathologistes a effectuer des évaluations comparables.

LABORATORY HANDBOOK OF MEDICAL MYCOLOGY by Michael R. McGINNIS,
662Ep ssuilin, 68" ,ahard cover, 91980. AcademicePress a N.Y-

"Owing to the increased importance of fungi in medecine, there is
a pressing need to discuss important topics such as laboratory safety and
emergency procedures, quality control and modern concepts,... to assist
microbiologists in safety isolating and acurately identifying fungi of
medical importance.... The key to understanding any field of science ne-
cessitates a thorought understanding of its language..." (the author).
Thus, the author devotes its first chapter to the morphological terms
to be use in modern medical mycology for the description and the classi-
fiecarionpoltetve pLuncie while chapter 40(2331p.) ~and.chapter 9 (63 1p..)
describe the systems of classification, detail the methods of study and
comment the diagnoses of genera of, respectively the Ascomycetes, Coelo-
mycetes, Hyphomycetes and Zygomycetes (sic) in the former and the yeasts
in the later. Chapter 9 (50 p.) adds a'synopsis of the mycoses, descri-
bing and illustrating symptoms, etiology and therapeutics of each disease.
The other chapters of the book are descriptive of technics: laboratory
safety, handling clinical specimens, bioessay procedures, susceptibility
testing, quality control, equipment maintenance, culture collection and
culture media. Appendices provides with glossary, list of synonyms and
list of fungal records on man.

The book is no doubt interesting and useful, not only for the tech-
niques described, but as an effort of providing medical mycologists with
the developments in modern descriptive mycology, especially in the field
of the Hyphomycetes, after the First Kananaskis Conference of 1969.

Non obstant such evident value, the book presents several major
defects that myself at least, I do regret. In the general disposition of
the contents, the book might have been divided advantageously into two
parts, one part, taxonomic, including chapters 1, 4 and 5 (the techniques
excluded), chapter 9 and the appendices, the other part, technical, grou-
ping chapters 2 and 3 with chapters 6 to 10, and the culture methods from
chapters 4 and 5. That grouping might have avoided regrettable duplications
between chapter 1 and chapter 4 in text and illustration.

The illustration is, beside a few line drawings, essentialy made
of black and white microphotographs of fungi, 11 x 8.5 cm (half page) in
size, many of them being of good quality. Unfortunately, the illustration
isdefective in several ways. The magnification is not given. The numbering
of the photographs, composed of the chapter number and the figure number
within the chapter is complex and unconvenient to the reader. Furthermore
from a count of 219 numbered figures, 52 are composed of 2 or 3 separate
full-size photographs, not mounted in plate, but printed on distinct pages
most often and provided with a full legend under fig. -A which is said
"continued" under fig. -B and fig. -C. This constitutes another unconve-~
nience. The procedure is in most case unjustified. Either the photographs
composing a figure show the same fungus at low and high magnification, the
one at low magnification being generally of no use or no value; or the
photographs represent different fungi and could have been numbered and
legended separately. At the same, the duplication of the same or similar
pictures of the same fungus, illustrating here a term and further a gene-
ric concept might have been avoided. For instance, a picture of Drechs-
lera sp. fig. 1.1 is the inverted photograph fig. 4.46 of the same. Why
Trichothecium roseum needs to be shown at different focusing in figures
4.105A and 4.105B on p. 303 and 304? Why Syncephalastrum ramosum must

284

appear ‘in 5 figures (1.9A, 1.9B, 4.120A, 4.120B, 4.121) and on distinct
pages? Unacceptable is the fact that Fxophiala spinifera is not only
figured three times in figs.1.27, 4.13 and 4.50 but also duplicated in
fig. 4.49A, identical to; fig. 4.13, under the name Exophiala jeanselmei.
It will be also mentioned that amongst the 161 figs. illustrating chap-
ters 1 and 4, 50 figs. are of unidentified species of filamentous fungi.
Why, in such an handbook designed to be informative as much as exampla-
tive, the author choose to illustrate unidentified isolates rather than
properly identified fungal species of medical importance which are avai-
lable in pure culture in official culture collections? Why finally the
photographs are supplemented of such poor and inacurate line drawings?

Other defects are in the text. The author decided not to cite
the authors of the names of the fungi in the text, but did cite them,
in Appendic C, only for the correct names of which he listed a synonym
but not for many other names used in the text. The reader will have to
consult other sources to distinguish for instance Blastomyces dermatiti-
dis (conidial Ajellomyces dermatitidis) from Wangiella (Hormiscium,
Phialophora, Exophiala) dermatitidis.

At the exception of the references inserted in the comments fol-
lowing the generic descriptions of the fungi, references to the litera-
ture are totaly absent from the general text, like are the references
to published sources for the 77 culture media cited. The reader need to
be informed on the source of every statement, term, definition, classi-
fication, key, technique or formila other than the author's personal

ones. : : ; ; i
Dealing with terminology in chapter 1, the author restrict the

application of the term chlamydospore to those terminal inflated thick-
walled cells which are able to germinate and be reproductive and pro-
poses the use of the term vesicle for the not germinating terminal
chlamydospores as well for the intercallary ones. This proposition should
be disrecommanded, the term vesicle being currently designating the
ampulliform tip of the Aspergillus conidiophore stipe. Further the author
illustratesthe term favic chandelier, in use in medical mycology, by a
photograph of the appressorial hyphae in Botrytis cinerea. AS soon as
1870, J. KLEIN introduced the term appressorium to designate that type
of hypha differentiated for the attachment of the fungus. There is no
reason for abandoning that term.
A-final) remark as néededvto infimum the, author s statement ti la—
gellate cells are produced only by the chytrids" on page 1 to 2.
Published by Academic Press, this handbook takes benefit of a
high quality typographic printing and binding, as it will surely be
in the hands of many mycologists in medical centers and laboratories.

FUNGAL, BIOTECHNOLOGY, by J.&. SMITH, D:R. BERRY & BS KRIST IANSEN,
x #308 ips, Ul. .68 ff hard cover, 419808 The British*Mycolozical
Society, Symposium series n° 3. Academic Press, 111 Fifth Avenue,

NewYork, NY10003..°) Price $326.50:

This book is the proceedings of a joint symposium of the British
Mycological Society and the Society of Chemical Insdustry, held in Glas-
cow in September 1978. The recent developments of the fungal biotechno-
logy has incited more research on the uses of fungi in the industrial
processes. The book is a selection of topics related to the inoculation
the growth and the exploitation of the fungi through the operation of
liquid and solid state fermenters. The parameters of fungal growth, from
the preparation of the inoculum, the genetical hazards, the rheological
conditions to the reactions to the type of fermenter and the nature of
the developed particles, are largely investigated and discussed for their

BOD

interference in the design of the fermenters. The production of edible
fungi is also considered. The potentialities of fungal biotechnology
are reviewed and the achievements in different regions of the world
also described. Although the repeated orthographic errors in the names

of the fungi, this book remains a very interesting account to applied
mycology.

FUNGAL SAPROPHYTISM by Harry J. HUDSON, Studies in Biology n° 32,

2d ed., 76 p., ill., paper back, 1980. Edward Arnold Publ. see
Bedford Square, London WC1B 3DQ. Price £ 2.25.

It is good to realize the author's ability to reach so many goals
through these modest 70 pages: enhancing curiosity and interest for the
fungi, considering many aspects of the fungal ecology, guiding the stu-
dent on the field and in the laboratory and demonstrating the practical
interest of the fungi in industry and human life. Some titles: wood de-
cay, blue stain, basidiomycetes and microfungi in litter, chitinolytic
fungi, lignine decomposition, pyrophilous fungi, coprophilous fungi,
moisture requirements, osmophilous fungi, storage fungi, xerophytes, aw
quatic fungi, water pollution, thermophilous fungi, composts, fermented
foods, antibiotics, vitamins. An exciting introduction to mycology,
through ecology.

FUNGI, MAN AND HIS ENVIRONMENT, by R.C. COOKE, 144 p., 111.,
paperaback ./1L9/7/7. Longman. Group, Ltd, London. Price £3.45.

"T have attempted to write a book about fungi rather than a text-
book on fungi''. The author indeed realizes a similar approach to that
of Houston, but with many more information and considering also the
parasitic fungi, appealing the awareness of people on the danger and
value of the fungi in the present and future world. Fror® the pests and
mycoses to their control, from the degadation of organic matter to the
synthesis of protein and the cultivation of mushrooms for foods and drugs,
the author clearly demonstrates the multiple both destructive and elabo-
rative potentialities of the fungi, in which, finally, man is concerned.
With science and philosophy, he suscites the desire of reading more
on fungi.

DIE BLATTERPILZE (AGARICACEAE) DEUTSCHLANDS UND DER ANGRENZENDEN
LANDER BESONDERS OESTERREICHS UND DER SCHWEIZ, par Adalbert RICKEN
ShUy +4462 p.,el22 pl. cols, im 8 ¢ relies heipzi¢gyl 9152 Reimprime
par Dr. Massimo CANDUSSO, Viale Europa 5, 21047 Saronno (Varese)
italia. Disponible. dans lL"allemand orzreinal: (Lit. 98,000:— net)

om traduLt envitalven par Dr. Lazzari (Lit., 82,000)— net):

La réimpression de 1l'oeuvre mycologique originale et unique de
Adabert RICKEN (1851-1921) est un événement heureux. Publiée et distri-
buée par livraisons successives de 32 pages et 8 planches colorées de
1910 4 1915, l'oeuvre de RICKEN fut trés bien accueillie par le monde
mycologique de 1'époque d'aprés guerre . En effet 1l'oeuvre est le bilan
de quarante année de récoltes mycologiques (1870-1910) en Allemagne et
régions limitrophes et portant sur plus de 1500 espéces. L'ouvrage pré-
sente dans un ordre systématique la description de 14£2 espéces d'acarics
sur la base d'observations précises tant microscopiques que macroscopi-
ques, avec mention de l'habitat et, pour 662 espéces, une illustration

286

en couleurs reproduisant les peintures a l'huile que réalisa 1l'auteur
sur le frais. La nomenclature est traditionelle mais étayée des dates
de publication. La derniére livraison comporta outre 1l'introduction et
les index alphabétiques trois clés dichotomiques des genres, fort bien
construites et basées sur les caractéres les plus évidents.

La réimpression entreprise par 1'Imprimeur mycologue, Dr. Massimo
CANDUSSO est particuliérement réussie. Le texte est reproduit de maniére
réguliére sur du papier de qualité. Les planches en couleurs, éditées
sur un carton plus fort comme les originales, reproduisent les tons
les plus stables des différents exemplaires originaux comparés. La re-
liure de type ancien est préservée dans une gaine. La qualité de cette
réimpression mérite les encouragements du plus grands nombre de myco-
logues.

ICONOGRAPHIA MYCOLOGICA par Giacomo BRESADOLA, avec J.B. Traverso,
L. Fenaroli. CG. Catoni® & J.B. Trenervediteurs, 26evols. 256.'pe.,
1250 pl. col., Milan 1927-1933. Réimpression par Dr. Massimo
CANDUSSO, Gruppo Micologico G. Bresadola, Viale Europa 5,

210477 Saronnes (Varese) Ltalia, in 5.volse,)l2b0rph.. col were lie,
Sept. 1981-Déc. 1982. Prix souscription Lit. 100.000 before
September 19895 Lit-100 S000 vamsrecumdese vols elie? sandy ale
solde Lit. 60-85.000 au recu du 4e vol.

L'examen de la planche proposée par 1'imprimeur CANDUSSO au Ssous-
cripteur semble présenter toutes les garanties de qualité que l'on peut
espérer, voire exiger, d'une telle réimpression. L'ouvrage de Bresadola
est une oeuvre majeure dans la connaissance des champignons. La réimpres-—
sion ne sera réalisée que si la souscription est suffisante. On ne peut
que recommander aux mycologues et aux institutions de science mycologique
l'acquisition d'un ouvrage que les générations futures de mycologues a
travers le monde ne pourront ne pas connaitre ni ne pas consulter.

Also nrecertved:

LABORATORY GUIDE FOR IDENTIFICATION OF PLANT PATHOGENIC BACTERIA
par N.W. SCHAAD , 72 p. 4 pl. col., mimeographed,4° in folder, 1980.
The American Phytopathological Society, St Paul, USA. PrixUS $10.-

MYCOTAXON

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MYCOTAXON

Roiemxitl, NO. 2, pp. 209-350 July-September 1981

THE GENUS PILOPHORUS

H. MARTIN JAHNS

Botantsches Institut der Universitat Frankfurt, Germany

Summary

All species of Ptlophorus are examined and four species are ex-
cluded from the genus. The remaining 10 species are described and
their geographical distribution is demonstrated on maps. Character-
istics used for the separation of species are discussed and an
artificial key is given.

Introduction

Formerly the genus Ptlophorus was placed in the Clado-
niaceae (Zahlbruckner, 1926), while today the genus is
thought to be related to the genus Stereocaulon (Jahns,
1970a; Henssen and Jahns, 1973). In an earlier paper I
attempted to delimit some species within the genus (Jahns,
1970b), but the choice of the species investigated at that
time was arbitrary, resulting from the context of my onto-
genetic work, so that no claim for completeness could be
made. Furthermore, the investigations of most species were
based on relatively limited material, so that conclusions
about variation of characteristics and geographical dis-
tribution were not reliable. For these reasons it seemed
desirable to complete the earlier observations and to ex-
tend the work to all species described by other authors
and at the same time to include more specimens. This is
the aim of the present work. Most of the specimens avail-
able from the major herbaria were used in this investi-
gation (more than 1000 specimens). A complete list of
their geographical distribution is not included as it
would ask to much space. For all practical purposes it
seemed to be sufficient to characterize the occurring
species, to mention their most important synonyms, to
distinguish them from each other and to demonstrate their
Gistribution on maps. The exact places of origin are
given for specimens found outside the main area of dis-
tribution. For a complete list of literature and synonyms
see Zahnlbruckner (1927b, 1939). ;

Material and methods

The material investigated was received on loan from the following
herbaria: Herbarium University of Michigan, Ann Arbor (MICH); Herba-
rium of the University of Georgia, Athens (GA); University of Texas
Herbarium, Austin (TEX); Universitetets Botaniske Museum, Bergen
(BG); Herbarium University of California, Berkeley (UC); Botanisches

290

Museum, Berlin (B); Botanisches Institut, Bern (BERN); Museum of Natu-
ral History, Budapest (BP); Farlow Herbarium of Cryptogamic Botany,
Harvard University, Cambridge (FH); Commonwealth Scientific & Indu-
Sstrial Research Organization, Division of Plant Industry, Canberra
(CANB) ; Chicago Natural History Museum, Chicago (F); University of
Alaska, College (ALA); Botanical Museum and Herbarium, Copenhagen (C);
National Botanic Gardens, Dublin (DUB); Forschungsinstitut und Natur-
museum Senckenberg, Frankfurt (FR); Conservatoire et Jardin botani-
ques, Geneva (G); Botaniska Tradgard, Gdteborg (GB); Department of
Botany, Dalhousie University, Halifax (DAL); Botanical Museum, Helsin-
ki (H); Botanical Institute of Hiroshima University, Hiroshima (HIRI) ;
Botanisches Institut der Universitat, Kiel (KIEL); Rijksherbarium,
Leiden (L); Herbarium of the Department of Systematics and Plant Geo-
graphy of the Botanical Institute of the Academy of Sciences of the
USSR, Leningrad (LE); Institute of Botany, Faculty of Science, Lisboa
(LISU) ; British Museum, Natural History, London (BM); Botanical
Museum and Herbarium, Lund (LD); Herbarium of the University of Wis-
consin, Madison (WIS); Botanisches Institut, Marburg (MB); Herbarium
A. Henssen, Marburg (Hsn); Botanische Staatssammlung, Mtinchen (M);
Herbarium of Yale University, Osborn Botanical Laboratory, New Haven
(YU) ; New York Botanical Garden, New York (NY); Hattori Botanical La-
boratory, Nichinan (NICH); Botanisk Museum, Oslo (0); National Museum
of Canada, Natural History Branch, Ottawa (CAN); Oulun Museo, Oulu
(OULU); Oxto Botanico, Padova (PAD); Muséum National d'istoire Natu-
relle, Laboratoire de Cryptogamie, Paris (PC); Botanical Department
of the National Museum, Praha (PR); Division of Botany, National Her-
barium, Pretoria (PRE); University of Washington, Herbarium, Botany
Department, Seattle (WZU); Botanical Department, Naturhistoriska
Riksmuseum, Stockholm (S); Botanical Department of Tromsoe Museum,
Tromsoe (TROM); Botanical Department, Museum of the Royal Norwegian
Society for Science and Letters, Trondheim (TRH); Botanical Institute
of the University, Turku (TUR); Institute of Systematic Botany, Uni-
versity of Uppsala, Uppsala (UPS); US National Museum, Department of
Botany, Washington (US); Naturhistorisches Museum, Wien (W); Botani-
scher Garten und Museum der Universitat, Ztirich (Z).

The specimens were mainly investigated with a stereo-microscope
and some typical material was cut with a freeze-microtome.

The genus Pilophorus and its subdivision

In my earlier Work I explained in detail why in my
Opinion the correct name of the genus and the correct
epithet are “Pilophorue .Th.Fr.". Further discussion of
this question followed from Culberson (1970), who favored
the name " Ptlophoron (Tuck.)Th.Fr." and from Hawksworth,
James and Laundon (1972), who agreed with my interpreta-
tion. Closely related to this problem is the question of
the type species, which in my opinion should remain P.
robustus Tn. Fr.4 (Jahns? 1970p).

In the last hundred years not only has the number of
species subsummized under this genus fluctuated conside-
rably, but attempts have also been made to subdivide the
genus. An extreme proposal was the classification of
Tuckerman (1882) who considered P. cereolus to be the only
species of the genus - designating all other species
earlier described as subspecies of P. cereolus.

Stitzenberger (1862) included the genus Helocarpon Th.Fr.

Pasi

as a subgenus in the genus Ptlophorus. He referred implicit-
ly to the description of Th. Fries (1861), who mentions
for Helocarpon crasstpes short black, stalked fruitbodies.
Fries himself obviously did not consider a relation bet-
ween the two genera described - Ptlophorus and dHelocarpon -
but later declared Helocarpon to be a synonym for Leetdea
fits Fries,) 10/74). Having examined the type material of
Lectdea crasstpes, I can confirm that there is no anatomical
or morphological similarity with any species of Ptlophorus.

A different subdivision of the genus was proposed by
Satd6 (1940). When giving the description of the new spe-
cies P. ntgrtcaule he believed there were principal diffe-
rences between this species and all other representatives
of the genus. He therefore suggested a division of the
genus into a section Mgrtcaule, including P. ntgricaule,
and a section E£uptlophorus, including all remaining species.
It was explained earlier (Jahns, 1970a) that this subdi-
vision cannot be accepted, as Sat6 based his description
Simones new Section on a, far too rigid partition of, tissues
pHowoOn pagmentation of the stalk tissue which, with varia-
ble intensity, can be found in other species too.

On the whole there seems to be no justification for any
subdivision of the genus.

The importance of different characteristics
for the separation of species

For the separation of the species the same criteria
were used as described in the first paper dealing with
part of the genus (Jahns, 1970b).. However, it is now pos-
Sible to give a more accurate definition of the range of
variation and to judge the value of the different charac-
berigstics more rigidly.

Thallus horizontalis

The thallus horizontalis is always crustose and is ge-
nerally grey or grey-green in color when dry. It consists
Sreipawutve granules (fig. 1-4) or of flat or inflated
scales (fig. 8). Small granules or scales may be pressed
together, forming distinct areolae (fig. 7). In some spe-
cies, especially in older material, the thallus horizon-
talis may be almost absent. The shape of the horizontal
thallus is clearly specific and relatively constant for
each species. Therefore it may be used as a distinguishing
eaeracveristic, but on its own it is not sufficient for
the separation of the species. Principally the external
morphology may be characteristic for each species but a
certain degree of variation in shape is induced by factors
like substrate, etc. Thus intermediate forms and transi-
tions between the habits of different species may occur.
Furthermore, variations in shape of the horizontal thallus
among different species are difficult to explain, so that
this characteristic is of restricted value for use in
ertificial keys.

Zoe

Thallus verticalis and growth form

The thallus verticalis of Ptlophorus is a pseudopodetium
similar to the one found in the genus Stereocauton. The po-
sition of the apothecia is always terminal. The pseudopo-
detia may be simple or branched. Their size varies within ~
certain limits but offers a useful “aid for the description
of the species. In nearly all species a few fertile speci-
mens occur where pseudopodetia are absent and the apothe-
cia are sessile on the thallus horizontalis. The habit of
the pseudopodetia offers one of the best means of distinc-
tion for taxonomic use. Most common are simple, unbranched
and pin-shaped pseudopodetia, which occur in P. cereolus, P.
strumaticus, P. nigricaule, P. fibula and P. ecurtulum (fig. 3,6,
7,8,10). If the pseudopodetia of these species bear more
than one fruitbody, an indistinct terminal branching may
be simulated. In contrast to these pin-shaped species,

P. robustus and P. actcularts have high, branched pseudopode-
tia which grow in dense clusters (fig. 1,9). An interme-
diate growth form between these two types is represented
by &. vegae, P. awasthianum and Py. clavatue.” Their pseudopode—
tia are of middle height and somewhat branched and, at
least in P. vegae and P. awasthtanum, may grow in more or
less distinct clusters Gligce 2,11). Quite tyoica nm toceau
species is their manner of branching, which is umbellate
in P. robustus (fig .9), and Torked=in Ps acreular1s, =) = veqac
and P. awasthtanum.

The pseudopodetia are usually covered with an algal
layer which is not continuous and often resembles the gra-
nules and areolae of the thallus horizontalis. Older
stalks may be partially bare. In P. ntgrtcaule the stalk is
covered with conspicuous peltate scales, which are closely
pressed together (fig. 8,17).

Soredia

In all species of Ptlophorus the pseudopodetia are cover-
ed with an algal layer which is never continuous but com-
posed of more or less dispersed areolae. A true cortex is
only poorly developed and usually completely absent, so
that the pseudopodetia in some species have a granular
appearance. True soredia only occur in P. cereolus, where
they give a distanet powdery Took to the stalk™ (1727) 5).

Cephalodia

Cephalodia are characteristic for all species of the
genus Ptlophorus. However, two species which have been in-
cluded in this genus (P. conglomeratum and P. stauffert )
lack cephalodia. This, among other reasons, justifies in
my opinion the exclusion of these species from the genus
Ptlophorus.

The cephalodia of Ptlophorus are sacculate and exhibit no
special wall structure. In the three tallest species, P.
robustus, P. actcularts and P. vegae, some cephalodia occur on
the pseudopodetia (fig. 1,9,11), but in all other species
they are only to be found: on the thallus horizontalis.
Their shape is irregular, the surface smooth or granular,
and they are light brown to deep brown-black in color.

The cephalodia either contain algae of the Nostoe or of
the Sttgonema type. In the species investigated I found Stt-
Goneia An Pe strumaticus, P. fibula, P: curtulunm -and P.-ntigrtcaule,
while in P. robustus and P. awasthtanum I found only Wostoe.
In P. aeteularts I observed only Wostoe, but Kurokawa and
phibuichi (1970). found Stigorema an this’ species. This con-
Gradi ction may not result from a misobservation, because
in P. cereolus, P. clavatus and P. vegae I found some cephalo-
dia with Sttgonema while others contained Wostoc . However,
in general, all cephalodia of one specimen contain the
same alga. Exceptions are rare. Cephalodia of P. vegae con-
tain Wostoe and Sttgonema and possibly algae of the Gloeo-
capsa type as well.

Nostoe bearing cephalodia often have a habit distinct
from those with Sttgonema. They are always characterized by
a smooth surface, while cephalodia with Sttgonema as phyco-
bionts show a granular surface, at least in their younger
stages of development. This phenomenon is explained by the
development of the cephalodium which was described else-
where (Jahns, 1972).

It is not impossible that from those species which seem
to contain only one algal species in their cephalodia,
specimens with other phycobionts will be found in the fu-
ture. The occurrence of specific blue-green algae in the
cephalodia is therefore not a useful taxonomic characte-
sou en wal one

Pycnidia, ascospores, fruitbody development

Pyenidia occur abundantly in all species of the genus
Ptlophorus. They are either located on sterile pseudopodetia
Oreonnine thallus norizontealis.= The conidiophores: are un-
branched and the conidia sickle-shaped. The size of the
ecoriidia does not differ significantly in the different
Species, so that this characteristic is unimportant for
the taxonomy of the genus. The same holds for the ellip-
Soid ascospores.

The development of the fruitbodies is very uniform too
(Jahns, 1970a). Some specimens of P. clavatus form an ex-
ception. Numerous young apothecial primordia are located
maebundles®on short stalks) (fie. 4)0 In the further
course of development the primordia fuse and, with the
growth of the generative tissue, form the typical elongat-
ed fruitbody of the species (fig. 5,16).

Anatomy and morphology of the apothecium

The outer shape and inner structure of the apothecium
belong to the best characteristics for the separation of
the species. In most species the mature apothecia are al-
most globose and quite often several apothecia are late-
mallyitused) (fies 19510). An exception as “the, fruitbody
of P. elavatus, where the hymenium extends a considerable
distance down the pseudopodetial stalk so that the apothe-
cium assumes a triangular outline in longitudinal section
rine 5 :16)e. SmallerGvariations. of themspherical structure
are found in P. aqwasthtanwn and P. ntgrteaule, which'both tend
towards slightly triangular-shaped fruitbodies (fig. 2,8).

294

However, it is possible that the samples of P. awasthtanum
that I. examined were not fully developed and mature.

A longitudinal section reveals that the spherical shape
of thesavpothecia 1s, noteunitrormly Lormed.wApariet yona., cue
globose apothecia, fruitbodies may be flattened with a
margin of the hymenium that may hang down for a considera-
ble distance before curving upwards again to fuse with the
Sia Tie es Po

Anatomically, three major characteristics are of im-
portance: the boundary-texture, the pigment-boundary and
the columella. The boundary between the vegetative tissue
and-ithe generative tissue of the fruitbody,) which is
always easily recognized, may follow different patterns.
It forms a thin horizontal zone in the lower periphery of
thé. fruitbody animost species! (rigk 1) whereas int ee:
robustus and P. struwnmaticus the vegetative tissue extends
upwards into the generative tissue of the apothecium
(fig. 12) forming a cone-like columella.

The boundary between the vegetative and generative tis-
sues may be emphasized by two conspicuous structures. In
the transition zone the generative tissue may form a boun-
dary-texture built from closely interwoven, short-celled
hyphae (fig. 13). Often the junction of vegetative and
generative tissue is characterized by a deposit of black-
brown pigment, (fic 15), which is »referred to,aseche
pigment-boundary. The pigment-boundary bears little rela-
tionship to the boundary texture andmay or may not be
superimposed. Black pigment is often concentrated in the
subhymenial layers (fig. 14,15) as well as in the pseudo-
podetial tissue (for example in P. ntgricaule, P. vegae and.
P. strumaticus).

The columella, the boundary-texture and the pigment-
boundary are characteristics which help to separate the
species very accurately. These characteristics may occur
SiIngely or combamed.

Secondary products

Small amounts of several specimens were investigated by
TLC. The results obtained agree in general with those pub-
lished by Krog (1968) and Kurokawa and Shibuichi (1970).
Alls species contain atrancrineand zeorin. Stictic acidiwas
found in P. ntgrtcaule and P. vegae. Furthermore, Kurokawa
and shibttachi found, caperatic acid: in 2. elavatus, sand scon-
SsticticVacid’ in’ P. ntgrieaule cas’ well as* six other non-
identified substances in different species of the genus.
These observations could not be substantiated in detail.

In this study theschemical characteristics will now be
considered for taxonomy, as obviously only those substan-
eés that are common to all species occur in sufficient
amounts. In order to avoid seriously damaging the usually
limited material of the herbarial specimens it is some-
times impossible to demonstrate the presence even of atra-
norin and zeorin. Therefore, it is almost impossible to
obtain an exact and reliable picture of the distribution
of these substances with the methods described.

295

Substrate

The species of this genus grow exclusively on silicate
stone. As an exception single colonies may extend to rig-
idly pressed soil covering the stone. One specimen of P.
acteularits from USA, Washington (near Lake Cushman, Mason Co.,
S.East Olympias (F)) was found growing on dead wood. The nu-
merous other specimens from the same region suggest ex-
tremely favorable climatic conditions which enable the
lichen to colonize the unfamiliar substrate.

Distribution

The species of the genus Ptlophorus occur exclusively
in the northern hemisphere. P. conglomeratum, P. colensot and
P. stauffert described from Australia, New Zealand and New
Caledonia are no longer included in the genus Ptlophorus.
In the literature it is still maintained that P. actcularts
eccurs invAustraliavand Africa, (Cape of Good) Hope). These
statements refer exclusively to a note from Nylander
(1860) who claims to have seen the material in the Hooker
herbarium, but these specimens disappeared later. In the
year 1860 the separation of the species was not well de-
fined, so that it may well have been P. conglomeratum. In
any Case consider i this: information’ too doubtful to an-
elude in- the distribution maps.

The distribution maps of the individual species are
based almost exclusively on specimens which I have person-
ally investigated. Only ina few cases have I included
notes from other workers (Krog, 1968; Ammann and Ammann,
O69; Kurokawarsand Shibuichd,,. 1970), as Ll am certain that
the definition of species by these authors agrees with my
own views.

fer indavidual points’ inv the’ distribucion maps often
include material from a relatively large area. The number
of specimens examined is given for each point. However,
in many cases these numbers are almost certainly slightly
higher than the number of findings proper, as it was im-
possible always to recognize identical specimens from the
same gathering which had been distributed and preserved
in different herbaria. In those cases where a single
finding is represented by numerous specimens distributed
as an exsiccate it is only recorded as a single point.
whe exaeveplaces of origioneare indiecated@in détaLny only
when either very few such places are known or when those
places are outside the main area of distribution. Places
which are listed in the description of the species are
marked by a point, others not individually listed are
marked by a square on the maps. Findings taken from other
authors are marked by a triangle.

In general the following large-scale distribution was
established: One group of species ( P. actcularts, P. ntgrt-
caule, P. elavatus) occurs around the northern Pacific Ocean.
In the same area two more species are found which, how-
ever, are restricted to only part of this area (P. curtulum
in China and Japan, P. vegae in Alaska and the Bering
Strait area). One species (P. awasthitanum ) is restricted
to the Himalayas and another (P. ftbula ) to the eastern

parts of the USA. One species ( P.. strumattcus) occurs only

those parts of Europe which have an oceanic climate.

Very typical is the distribution of P. robustus, which is
found circumpolarly in the most northern areas and in
high mountains. P. cereolus shows the most irregular di-
stribuuion. [hie, species as most abundante ire ocandinavia
but occurs in single specimens in middle and central
Kurope, if Siberia and an the eastern part of North Ame—
rVCa.

Artificial key to the species

la
1b

3b
4a
Ab
5a

8b
9a

10a

Lichen with pseudopodetia and developed apothecia 2
Lichen. without pseudopodetia or pseudopodetia without apo-
thecia, sometimes with pycnidia or black primordia of apo-
thecia. 15
Thallus horizontalis and pseudopodetia clearly sorediate.
Pseudopodetia pin-shaped, unbranched, at most 1 cm high.
Apothecia without columella (fig. 3). P. cereolus
Surface not sorediate, at the most slightly granulose. 3
Pseudopodetia with continuous white cortex. Ecorticate

parts of the pseudopodetia and their central part always
black. Pseudopodetia 0.5=-1.5 cm high, usually dichoto-—

mously branched. Large cephalodia on short branches. Apo-
thecia unknown. Alaska, Bering Strait. (fig. 11) P. vegae

Lichen with other characteristics. 4
Pseudopodetia smaller than 5 mm. 5
Pseudopodetia higher than 5 mm. it

Apothecia cylindrical or triangular in longitudinal section,
at least twice as high as broad (fig. 16). Pseudopodetia
pin-shaped, unbranched. Apothecia often slightly bifurcate

atethe apices (higs.5) . P. elavatus
Apothecia globose, sometimes several globose apothecia
are fused. 6

Pseudopodetia very thin (0.2-1.0 mm broad, 5 mm high),

more or less branched, partly ecorticate and black (fig. 2).
Himalayas. P. awasthtanum
Pseudopodetia broader, pin-shaped, unbranched. q;
Scales of the thallus horizontalis and of the pseudopo-

detial cortex peltate (fig. 17), white, pressed together,

with an inflated appearance. Inner parts of the pseudo-
podetium with dark pigment. Apothecia with boundary-texture,
globose or sometimes pointed (fig. 8). P. ntgrtcaule
Lichen with other characteristics. 8
Thallus horizontalis formed by granules or minute scales.
Granules or scales fused into broader, closely arranged
areolae (fig. 6,7). £
Granules or scales of the thallus horizontalis scattered. 10
Apothecia always with boundary-texture and pigment-—boun-

dary. Pseudopodetia usually not higher than 1 mm (at

most 2 mm) (fig. 7). Eastern North America. P. ftbula
Apothecia always with boundary-texture, only very old
specimens with pigment-boundary. Pseudopodetia 1-3 mm

high (fig. 6). Japan and China. P. curtulum
Apothecia without columella, with boundary-texture and

in old specimens with pigment-boundary (fig. 6). Japan

and China. P. curtulumn

10b

lia

i aliss
12a

12b
13a

13b
14a

14b

15a
i Se
16a

16b
17a

17b
18a

18b

19a

19b

20a

20b
2la

21b
22a

22b

Apothecia with columella, without boundary-texture and
pigment-boundary (fig. 10,12). Europe. P. strumattcus
Apothecia cylindrical or triangular in longitudinal
section, at least twice as high as broad (fig. 16). Pseu-
dopodetia pin-shaped, unbranched. Apothecia often slight-
ly bifurcate at the apices (fig. 5). Peeelavacus
Lichen with other characteristics.

Scales of the thallus horizontalis and of the pseudopo-

.detial cortex peltate (fig. 17), white, pressed together,

with an inflated appearance. Inner parts of the pseudopo-
detium with dark pigment. Apothecia with boundary- texture,
globose or sometimes pointed (fig. 8). P. ntgrtcaule
Lichen with other characteristics.

Pseudopodetia very thin (0.2-1.0 mm broad, 5-15 mm high),
more or less branched, partly ecorticate and black

(fi¢. 2). Himalayas. P. awasthtanum
Lichen with other characteristics.

Pseudopodetia umbellately branched towards the apices.
Apothecia numerous, often fused at the margins, with

large columella (fig. 9,12). P. robustus
Pseudopodetia dichotomously branched, in combed tufts,
smaller plants unbranched. Apothecia without columella
‘Getkeps Hays P. acteularts
Pseudopodetia with pycnidia or with primordia of

apothecia present.

Only thallus horizontalis present.

Thallus horizontalis and pseudopodetia sorediate. Pseu-
dopodetia pin-like, often broadest in the middle

Gale ENS . P. cereolus
Lichen not sorediate.

Pseudopodetia very thin (0.2-1.0 mm broad, 5-15 mm high),
more or less branched, partly ecorticate and black

(fig. 2). Himalayas. P. awasthtanum
Pseudopodetia broader.

Pseudopodetia with continuous white cortex. Ecorticate
parts of the pseudopodetia and their central part always
black. Pseudopodetia 0O.5-1.5 cm high, usually dichotomous-
ly branched. Large cephalodia on short branches. Apothecia
unknown (fig. 11). Alaska, Bering Strait. P. vegae
Lichen with other characteristics.

Thallus horizontalis persistent, formed by closely
connected areolae. Sterile pseudopodetia 1-2 mm high,
terminated by a pointed pycnidia. P. clavatus
Thallus horizontalis formed by scattered granules,
evanescent in older specimens.

Pseudopodetia at least indistinctly umbellately branched
in their upper part. With numerous pycnidia or primordia

of apothecia. P. robustus
Pseudopodetia slender, unbranched or dichotomously
branched. P. actcularts
Thallus horizontalis not sorediate, sometimes granular.
Thallus horizontalis sorediate. P. cereolus

Thallus horizontalis formed by white, peltate,

inflated scales or granules. Japan, west coast of

North America. P. ntgricaule
Lichen with other characteristics.

155

16
4)

ue)

Zod.

Boo

23a Thallus horizontalis persistent, formed by closely
connected areolae. 24
23b Thallus horizontalis persistent or evanescent, formed
by more or less scattered granules or scales. Determi-
nation only from distribution (see maps). P. robustus,
P. strumaticus, P. acteularts, P. vegae, P. awasthtanum,
P. curtulum
24a Areolae thin, not distinctly formed by individual

granules. P. clavatus
246 Areolae thick, composed of small granules. ao
25a Distribution in Japan and China. P. curtulum

25b Distribution in-the eastern North America. P. ftbula
Description of the species

Pilophorus acicularis (Ach.) Th.Fr., De Stereocaultis et Ptlophoris
Comm. + “41° 41857);

Baeomyces actcularts Ach., Meth. Lich.: 328 (1803). Type collection:
North America, Hall (UPS, isotype).

Cenomyce acteularts Ach., Lich. Univ.: 567 (1810).

Thallus horizontalis small granular, evanescent in older thalli,
grey when dry. Pseudopodetia (0.5-) 3 cm high, 1 mm diameter. Mostly
Simple or dichotomously branched, stalks curved so as to appear as if
combed; rarely erect, pin-like and 1 cm high (fig. 1). In some speci-
mens richly branched in the upper part of the pseudopodetia, thus

Ne
‘
os
70
ix
ALB.
,

‘a
wva'? WbED:
wpe

cu)

fig. 1 P. aeteularis (ce - cephalodium)

Z99

faintly resembling P. robustus. Internally the pseudopodetia are solid
when young, becoming hollow with age, composed of long, thin, strongly
gelatinized hyphae with narrow lumina (0.5 um). The lower part of
older pseudopodetia becomes blackened internally. Algal layer not
continuous, in more or less distinct granules, entirely absent from
some parts of the surface.

Pycnidia occur in the tips of small sterile pseudopodetia or in
the tips of small lateral branches of older pseudopodetia. Conidio-
phores 30 pm long, unbranched, with terminal sickle-shaped conidia,
6exoi> pm.

Apothecia abundant, one or several on the apices of the pseudopo-
detia, black, hemispherical or indistinctly triangular, up to 1.5 mm
diameter. There is no columella.

Hymenium up to 240 pm high, two-thirds pigmented, lower part ste-
rile, consisting only of paraphyses. Excipulum absent. Asci eight-
spored. Spores rounded when young, becoming spindle-shaped when ma-
pure, )21.0-29.5 x 4°75-5.5 pine Generative tissue formed¥of closely
interwoven hyphae with short, broad cells with large lumina. The ge-
nerative tissue is pigmented black-brown, the color being most in-
tense below the paraphyses, becoming less so towards the stalk
region.

Phycobiont green, Pleurococcus type, cells c. 9 pam diameter.
Cephalodia on the thallus horizontalis are brown and hemisphe-
rical and contain Wostoc. Kurokawa and Shibuichi (1970) found cepha-
lodia containing Sttgonema. Small cephalodia also occur on the pseu-

dopodetia (fig. 1).
Growing on silicate stone, seldom on decaying wood (one specimen).

Remarks

The correct citation for the species is P. actcularts
Phcony yin. Fre (105 /p. 4ooe ands not P..aetcularis (Ach...)
Ny 105) ep. 90) sae explained e€arlier (Jahns, 1970b);

P. acteularts can be separated from the smaller species
by its tall pseudopodetia. It may be confused with P. ro-
bustus, especially in material from Alaska where both spe-
cies occur together. But usually the type of branching
is different (umbellate in P. robustus, dichotomous in P.
acteularts) and the lack of a columella in longitudinal
sections of the fruitbodies of P. acteularts always makes
distinction easy.

Th. Fries (1889, p.765) described P. actcularts var.
econjugens Th.Fr. from Vancouver Island, which is said to
resemble P. robustus. I have seen the original material
and can see no difference from numerous other specimens
which are not worthy of varietal status.

Distribution (map 1)

P. acteularts is probably the most abundant species of
the genus. Most specimens were found on the west coast
Of North: America, with a focal point in British ,Columbia
and Washington. The species is widely spread in Japan
too. In general this species seems to prefer an oceanic
climate without extremely low temperatures, at least in
comparison with other species of the genus. This assump-
tion is supported by the fact that this species is found
more southerly (44 Tindings in California) than all ‘other

listed M in the description of the species)

species and only two specimens were found in northern
Alaska where, for example, P. robustus and P. vegae occur
more often. East of the Rocky Mountains P. actecularts is
rare too. It is very difficult .to give any estimate on the
occurrence of the species in China. Although there are on-
ly three places known to me where the species has been
found, at least one gathering was large enough for the
distribution of numerous specimens as an exsiccate (Krypt.
Vind. Exs. 2842). Therefore, it may be suggested that P.
acteularts is not rare in China, but simply has not been
collected.

Places outside the main distribution area (map 1, marked
by aucirele):
Canada, Ontario, Belleville (CAN).

Pilophorus awasthianum Rads., Arch. Soc. Zool. Bot. Fenn. Vanamo
5:28-29 (1950). Type collection: India orientalis, Himalaya
orientalis, Darjeeling, 3500 m (H, holotype).

Thallus horizontalis persistent or evanescent, consisting of
loosely scattered granules, c. O.1 mm diameter, grey when dry. Pseu-
dopodetia simple or branched, small and very thin, O.5-1.5 cm high
and 0.2-1.0 mm broad. Pseudopodetia partly covered by an algal
layer consisting of small granules, but for the most part the stalks

301

fig. 2 P. awasthtanum (ce - cephalodium, p - pycnidium)

are bare. The ecorticated parts are blackened, the algal layer grey.
Internal part of pseudopodetia solid, formed by strongly gelatinized
hyphae.

Pycnidia apical on short pseudopodetia.

Apothecia apical on mature pseudopodetia, sometimes simple and
globose but mostly aggregated, consisting of several subglobose bo-
dies (fig. 2). Apothecia 1.5-2.0 mm in diameter. In mature specimens
apothecium and pseudopodetium are separated by a pigment-boundary, ~
while boundary-texture and columella are absent. Hymenium c. 100 pm
high, excipulum absent. Asci eight-spored, spores spindle-shaped,

c. 20 x 7 pm.

Phycobiont green, Pleurococcus type, cells c. 10 pam in diameter.

Cephalodia contain Nostoc, black, globose, sessile on the thallus
horizontalis, c. O.1 mm diameter.

Growing on silicate rock and earth.

Remarks

P. awasthtanum differs from all other species by the
small, branched and very thin pseudopodetia and by the
strongly aggregated apothecia. As there are only two

S02

known specimens so far, the range of Variatvoneot ithe
different characteristics is unknown.

Distribution (map 2)

The species is only known from the Himalayas. I have
seen the following specimens:
India, Darjeeling district, E. Himalayas, Awasthi, 1948 (H, holo-
type); India Sandakhpu, Awasthi, 1950. (UBS).

map 2 P. awasthtanum - distribution

Pilophorus cereolus (Ach.) TH.Fr., Lich. Scand. 1, 1: 55 (1871).

Ltehen cereolus Ach., Lich. Suec. Prodr.: 89 (1798); Meth. Lich.:
316 (1803). Type collection: Sweden, Facklelaf, E. Acharius
(H, holotype).

Thallus horizontalis persistent, consisting of loosely aggre-
gated, sorediate granules, c. O.1 mm diameter, grey-green when
dry. Pseudopodetia pin-like, usually up to 5 mm (occasionally up
to 1 cm) high, 1.5 mm broad, sorediate. Sterile stalks or those

503

fig. 3 P. cereolus (ce - cephalodium, p - pycnidium)

bearing only pycnidia are usually broadest in the middle and irregu-
larly curved (fig. 3). In some specimens, especially in the few Ame-
rican samples, the upper part of the pseudopodetium is slightly
branched, each branch bearing a pycnidium or a primordium of a fruit-
body. Internally the stalks are compact, composed of strongly gela-
tinized hyphae with narrow lumina. The phycobiont layer is granular
sorediate.

Pycnidia bottle-shaped, abundant, several often produced at the
apex of a pseudopodetium. Conidiophores c. 25 pm long, unbranched,
with terminal conidia. Conidia sickle-shaped, 5 x 1 pm.

Apothecia scarce, solitary or seldom two on one stalk, spherical,
black and terminal, up to 1.5 mm diameter. In longitudinal sections
the apothecium is spherical and without a columella. The boundary
between the generative and vegetative tissue remains horizontal with
the lower margin of the apothecium. In old apothecia a strongly co-
lored pigment-boundary is situated at this juncture. An indistinct
boundary-texture may sometimes be developed. Hymenium c. 80 pm high,
almost unpigmented. Excipulum absent. Asci eight-spored. Spores
rounded when young, becoming spindle-shaped when mature, 14.5-21 x
5.5-6.5 pm. The regions just below the paraphyses and in the pigment-
boundary are deep black-brown, towards the center of the apothecium
the color grades to pale brown.

Phycobiont green, Pleurococcus type, cells c. 9 pm diameter.

304

Cephalodia brown, clustered,-sessile on the thallus horizontalis,
c. 1-2 mm in diameter. Their surface is usually wrinkled. The cepha-
lodia contain Nostoe or sometimes Sttgonema.

Growing on silicate stone.

Remarks

P. cereolus is distinguished from all the other species
by the sorediate thallus horizontalis and pseudopodetia.
Without this characteristic fruiteng specimens Onn wcere—
olus could besmistaken for PP, strwunaticus, Pay tbula,specureu—
lum or P. ntgricaule, as all these species are of about the
same height and pin-like in appearance. The pseudopodetia
of P. vegae sometimes have a granular cortex, which could
be considered as sorediate, but the habit of the species
distanculy disiers. fromeP. cereolus (ing — jeond) 1.) An
other important characteristic for the distinction of the
small, pin-like species is the longitudinal section of
the fruitboodiese( tre. ei 5 en

Tuckerman, in his later works (e.g. Tuckerman, 1882,
p.235) considered all other species to be varieties of
P. cereolus. As a result, many subsequent authors have
attributed all species of the genus to this species. As
pointed out by James (1965, p.239)°"P. cereolus" as used

map 3 P. cereolus - world distribution (places listed @ and not
listed M in the description of the species)

305

by British authors refers mostly to P. strwmatreus and not
to the present species. In the same way "P. cereolus" as
used by many American authors refers to P. actcularts.
Rasdnen (1946, p.2) described P. cereolus var. ecephalodt—
ferve has, which he saidvas to be distinguished by the
smaller pseudopodetia and the abundant, markedly tuvercu-
late cephalodia. The syntype material of this variety was
distributed in Lich. Fenn. Exs. as no. 960. The samples
Pre an no Way diiferént from a multitude of other poorly-
developed specimens and should be considered merely as ju-
venile specimens which are not worthy of varietal status.

map 4 BP. cereolus - distribution in Europe (places listed @ and
not listed MM in the description of the species)

306

Distribution maemo)

P aeneolus US thesmost abundant Species sor sence eciie aya
Europe. Most samples were collected in Scandinavia where
sts occurrence overlaps with the distribution of P. robustus i
and P.tetmeatieus. However, P. .strumaricus: Usyrestricved) Ge |
the oceanic climate of the Norwegian and Swedish west |

coasts, and P. robustus occurs in the Norwegian highlands
and in the Arctic only, whereas P. cereolus is spread over |

on 15 20

reo)

7 bey) Ss

CGT sth ROT
map 5 P. cereolus - distribution in Scandinavia (places listed @
and not listed # in the description of the species)

DOr,

Scandinavia, including Finland. The absence of P. cereolus
in Great Britain suggests that this species, unlike P.
strumatteus, dislikes an oceanic climate. This agrees with
the observations made by Ammann and Ammann (1969).

Outside Scandinavia P. cereolus shows an interesting
distribution (maps 3-4). These rare but widely-spread
findings are listed further on. While a specimen from
northern Poland links up reasonably well with Scandinavia,
the occurrence in Siberia appears somewhat isolated. How-
ever, the material from Siberia is well developed and was
most certainly very abundant as samples of this material
can be found in many herbaria. A somewhat bigger group of
findings originates from the Carpates and the Tatra (Cze-
choslovakia), partly distributed as an exsiccate (Lojka,
rc. Huet Oo!)

In the Alps this lichen is apparently far less abun-
dant than might be expected from the older literature.
Although I have not seen all material mentioned in older
publications it seems that most samples belong to Stereo-
caulon, especially Stereocaulon ptleatum, which is small and
sorediate and looks somewhat similar to P. cereolus. Nume-
rous specimens originate from the Oberpfalz in southern
Germany. These and material from South Tyrol have been
distributed as exsiccatae (Arnold, Lichenes exs. 823,
2008), but apparently it has been found only in a few
places. The lichen seems to be absent from the rest of
Germany and the more western countries.

The literature does not give any clue to the distri-
bution of P. cereolus in North America as P. cereolus, P.
acteularts and P. ftbula were completely confused until re-
Cenoly. In particular, P. actcularis was designated as, \P.
cereolus in many cases. Krog (1968) remarks, that she has
not seen any authentic specimen of P. cereolus from wes-
tern North America. She mentions one single specimen
which she has not seen herself, but which from the de-
Scription could have been P. cereolus. I had the chance to
see this sample and it proved to be P. aectcularts too. We
May, therefore, conclude that P. cereolus does not occur
in the western parts of North America. In the eastern
part of the continent there are three places where true
P. cereolus has been found. One of which (Grand Marais)
was described by Fink (1910) who, however, thought it to
be identical with the P. cereolus described by Tuckerman
and which in fact was P. acteularis. The American material
is not fully developed and slightly different from the
Kuropean material. Nevertheless, it seems to be the same
species and its distribution in North America is linked
bo, the Mam vdistributionm area in Europe by findanes from
Newfoundland, Greenland and the Azores.

Places outside the main distribution area (map 3-5,

marked by a circle):

Canada, Newfoundland, upper Humber River (H); Canada, Newfoundland,
New Harbour, Trinity Bay (M); Canada, Minnesota, Grand Marais,

along the northern shore of Lake Superior (4 specimens MICH, 1 speci-
men US); Canada, Minnesota, Cook County, 8 miles SW of Grand Marais

308

on US Route 61, Cascade River State Park, along Cascade River (CAN);
USA, New York State, Adirondack Region, Chapel Pond, near St. Hu-
berts (MICH); Greenland, Diskofjord, Kuanersuit suvdlat (C); Azores,
Santa Maria, Pico Alto (James and Henssen unpublished) (MB); Germa-
ny, Oberpfalz, above Spielberg near Nabeck west of Schwandorf (11
specimens BM, C, H, KIEL, M, O, PR); Germany, Oberpfalz, Schutzen-
gelsteinbruch, Eldensteiner Forst (M); Austria, Tyrol, Silz, Intal
(2 specimens M); Italy, South Tyrol, Margola near Predazzo (25 spe-
cimens:B, BM, ‘BP, FH; H; LD, M, 0, S, W)? sitaly, South Tyrol, Set—
ween Bellamonte and Paneveggio (4 specimens BM, BP, M, W); CSSR,
Presov, 0. Szatala (BP; CSSR, alpes Dzurowa prope pagum Teplicska,
com. Lipto (W); CSSR, Slovakia, Nizke Tatry, in valle rivi Svidovy
(PR); CSSR, Slovakia, Nizke Tatry, in valle rivi Svarinsky (PR);
CSSR, in sylva Stefkofka prope Bresztova, com Arva (16 specimens

B, BG, “BPS iFH,. G) H,;) LD, 3M; 0, PR iS; Us (oN) eo Poland, sproperhabiau
(H); USSR, Dudinka, Yenisey, lat. bor. 69°35! (7 specimens FH, H,
OS US re |

Pilophorus clavatus Th.Fr.,Bot. Notiser, 1888: 214 (1888). Type
collection: Canada, British Columbia, Vancouver Island, Mt.
Mark, 1887, Macoun (UPS, holotype).

P. acteularts £. hallit Tack./Preoe,; Amt Acad ArtstSci. sl2-sl77 |
(1877) . Type collection: USA, Oregon, Hall (UPS, isotype). |

P. cereolus var. hallit (Tuck.) Tuck., Syn. N. American Lich.: 235
(1882).

Py hatltit (Tack.) -Vain.;. BotzeMag., Pokyoe35=. 59n (1021 |

P. japontecum Zahlbr. (nomen nudum), Catal. Lich. 4: 432 (1926);

Bou, Mage, “Lokyoc41: 387i, G1927 je.

fig. 4 P. elavatus, with young fruitbodies (ba - bush-like pri-
mordia of apothecia, ce - cephalodium)

SOS

Thallus horizontalis thin, granular persistent. Granules often
aggregated, forming scales. Granules minute, O.1 mm diameter, grey-
green when dry. Pseudopodetia up to 2 cm high, c. O.3 mm thick (fig. -
5). Internally the central area of the stalk is massive and consists
of thin, strongly gelatinized hyphae with thin lumina (0.5 pm). Al-
gal layer not continuous, formed by more or less indiscrete granules,
the stalk usually being naked just below the apothecium. Central
lower part of the stalks sometimes blackened.

Pycnidia bottle-shaped, apical on short pseudopodetia, about 1 mm
high, often bearing thalline scales which form a collar halfway up
the stalk. Conidiophores unbranched, 25 pm long, with terminal coni-
tay OO ox. O 5 At.

Apothecia black, occurring
Singly at the apex of each
pseudopodetium, often spread-
ing some distance down the
stalk. The apothecia are tri-
angular in longitudinal sec-
eLOM eid. 16), about. 2 W=5)
mm high, 1 mm broad at the
upper end, tapering to O.4 mm
at the base, often they are
more or less bifurcated at
the apices (fig. 5). Apothe-
cium sometimes formed by fu-
Sion of several primordia
growing on bush-like branched
pseudopodetia (fig. 4). No
columella, boundary-texture,
or pigment-boundary is formed
between the vegetative and
generative tissue. Hymenium
c. 200 pm high. Asci eight-
spored. Spores rounded when
young, becoming spindle-
Shaped when old, 23.5-26.5 x
5.0-6.0 pm. Hymenium green-
ish, and the generative tis-
sue uniformly pigmented, black-
brown.

Phycobiont green, Pleurococcus type, cells c. 8 pam diameter.

Cephalodia brown-black, irregular, lumpy, about O.5 mm diameter
with a velvety surface. Growing on the thallus horizontalis. They
contain Sttgonema and sometimes Nostoc.

Plants growing on silicate rock.

figs 59h. elavacus

Remarks

In most specimens the triangular outline of the apo-
Cheevalissurficienteto dietinguish Pa clavatussirom all
other species. P. eclavatus could only possibly be confused
with small specimens of P. aeteularts bearing slightly tri-
angular apothecia. In this case the two species may be
separated by their persistent or evanescent thallus hori-
zontalis respectively.

Tf was *explained, earlier (Jahns, #970b) thatr2. hall
is a synonym for P. elavatus and that the name P. clavatus

Den

Th.Pr. (1608) has pricrity at the species Jever over.)
halit4 . (Tuck.) Vain. (1921). PF. japontewn Zanibraelseaanomen
nudum. Zahlbruckner knew it to be a synonym of P. halltt
but nevertheless preferred his new name.

Distribution (map 6)

The distribution of -P.~clavatus 13) essentially 1denivical
with the occurrence of P. acteularts. Both species are found
on the coasts lot the northern Pactra Ocean, a sme ardcue
apparently being relatively more abundant than P. aeteula-
rts in Japan. i surprisingly Pwelavatus, nas siiou yeu bec
found in Northern Alaska but was present on the Asiatic
side of the Bering Strait. Kurokawa and Shibuichi (1970)
have examined material from Formosa and their observation
is included in map 6 (triangle), they also mention the
species in Korea but don't seem to have examined the ma-
terial themselves.

fry eye
ech MAA

map 6 P. clavatus - distribution ( M® places not listed in the
description of the species, aA specimens not examined)

Pilophorus curtulum Kurok. and Shib., Journ. Jap Bot. 45, 3: 78
(1970). Type collection: Mt. Ontake, Prov. Hida, Japan (TNS,
holotype; FH, O, isotypes).

Thallus horizontalis persistent, consisting of minute granules,

O.1 mm diameter, usually aggregated forming small squamules, 1.5 mm

————E—EeEE——————

ea.

4
S

fig. 6 P. curtulum (ce - cephalodium)

diameter (fig. 6). Sometimes granules scattered and hardly aggegat-
ed. Thallus grey-green when dry. Pseudopodetia pin-like, 1-3 mm
high, 0.5-1.0 mm broad. Mostly covered by a continuous algal layer
consisting of small granules. Only sometimes ecorticate. Internally
the stalks are compact, composed of strongly gelatinized hyphae.
Central part of older pseudopodetia colored by deposition of dark
pigment.

Pycnidia sessile on the thallus horizontalis.

Apothecia apical on the pseudopodetia, nearly always single, but
sometimes two apothecia developed from two primordia have partly
merged. Sometimes apothecia sessile on the thallus horizontalis.
Apothecia about 1 mm in diameter. In longitudinal sections the apo-
thecium is spherical and without columella. Pseudopodetium and apo-
thecium are separated by a boundary-texture which is formed in the
young primordium. In older specimens the boundary-texture is masked
by a pigment-boundary. Hymenium c. 150 pm high, almost unpigmented.
Excipulum absent, asci eight-spored. Spores rounded when young,
becoming spindle-shaped when mature, c. 7 x 22 pm.

Phycobiont green, Pleurococcus type.

Cephalodia contain Sttgonema, they form dark-brown flattened
discs between the squamules of the thallus horizontalis.

Growing on silicate rock.

lee

Remarks ,

This species resembles P. ftbula. The thallus horizonta-
lis of P. curtulum seems to be a bit thinner and less areo-
lated, and the pseudopodetium is somewhat higher and more
granulated than ain P.. frbula; but probably theyonLy ‘cood
reason for keeping the two species separated is their dis-
JUNCteGLStrrpution. Pe curtuium is only Known vomeapane ane
China, while P. ftbula is endemic to north-east America.

Distribution (map 7)

This lichen has been found in Japan several times. I
have seen samples from 10 places. From China I have seen
two samples:

Cleblieel, Vathewete, lens, creennel, (@)) 9 Claalinel, wAGheNeia, leenraevceulGl. 6 Ike imines
Waha prope pagum Yungning regione frigide temperata infra casulam
Maoniubi, alt. 3800-4030, Handel-Mazetti (Diar.Nr.1286) (W).

OS BAC:

map 7 P. curtulum - distribution (places listed e and not
listed ™ in the description of the species)

Pilophorus fibula (Tuck.) Th.Fr., De Stereocaults et Ptlophoris
Comm.: 42 (1857).

Stereocauton ftbula Tuck., Proc. Am. Acad. Arts Sci. 1: 238 (1847).
Type collection: USA, New Hampshire, White Mts., 1844, Tucker-
man (FH-Tuckerman, holotype).

Thallus horizontalis squamulose, squamules large, up to 2 mm dia-

el ee

meter, composed of small granules, persistent, grey-green. Pseudopo-
detia seldom exceeding 1 mm hight, sometimes totally absent so that
the apothecia are sessile (fig. 7). Stalk massive, composed of gela-
tinized thin hyphae with small lumina (0.5 pm). Algal layer more or
less areolate, not continuous.

Pycnidia single, in small diminutive pseudopodetia. Conidiophores
ce 30 pm long, unbranched, with terminal sickle-shaped conidia,

6 x 1 pm.

Apothecia black, hemispherical, 1 mm in diameter. A pigment-—boun-
dary and boundary-texture lies between the generative and vegetative
tissues. There is no columella. Hymenium up to 120 pm high, lower-
third pigmented. Lower edge of apothecium sterile, consisting only
of paraphyses. Excipulum absent. Asci eight-spored. Spores rounded
when young, becoming spindle-shaped when old, 17.0-23.0 x 5.5-6.5 pm.
A dark-brown pigmented layer occurs under the paraphyses, which be-
comes paler towards the center of the apothecium.

Phycobiont green, Pleurococeus type, cells c. 6 pm diameter.

Cephalodia contain Sttgonema, they form brown, Be Ee granular
discs, c. 1 mm diameter between the scales of the thallus horizon-
ee S ta ea ee

Growing on silicate stone.

fig. 7 P. fibula (ce - cephalodium)

Remarks

Since the first description of Stereocaulon ftbula by
Tuckerman in 1847 this specific epithet has tended to be
applied to widely differing small or depauperate speci-
mens of the genus. But P. strumattcus and P. cereolus differ
distinctly from P. ftbula by the occurrence of a columella

314

in the apothecium or of soredia respectively. Most diffi-
cult: to" distineulseh trom Py tbulatisw?. curren sea one
differences between the two species are only gradual. The
thallus norizontalis of 2, curtulio 2svusually bul noe always
less squamulose and less persistent. Moreover, only very
old apothecia of P. ecurtulum are separated from the stalk

by a pigment-boundary in addition to the boundary-texture
which is always present.

Distribution (nao fo
P. ftbula occurs only in north-east America and appears
to be restricted mainly to the area of the White Mountains.
Only two samples originate from the coast. I have seen ma-

terial from *the following places:

USA, New Hampshire, Mont. Alb. (BM, FH, M, MICH, W); USA, New Hamp-
shire, Roche entrance to Ilume, Franconia (F, US); USA, New Hamp-
shire, montibus Nord Anglio (PC); USA, New Hampshire, Warren (US);
USA, New York, rock on trail up Mt. MacDetyse near Lake Placid (MICH) ;
USA, New York, Adirondacks, Mt. Marcy (S); USA, New York, Adiron-
dacks, Indian Falls, near Mt. Marcy, near Lake Placid (FH); USA,
Maine, Mt. Desert Island (M, US).

map 8 P. ftbula - distribution ( e places listed in the description)

epke

Pilophorusenigricaule Sato, Journ. Jap. Bot. 16: 173)(1940) £1g.7.
Type collection: Japan, Honsyt, Mt. Tydkai (Tokyo, holotype).
Pr lopnorom cereolus (non Th.Fr.) Sat6, Journ... Jap; Bot. 9: 214° (1933).
Thallus horizontalis persistent, white or light grey, granular.
Granules c. 2 mm broad and c. 1 mm high, subglobose, aggregated or
scattered on the substrate. Most granules slightly peltate. Pseudopo-
detia pin-like, 1-6 mm high, 1 mm diameter. Internally the stalks
are compact, composed of strongly gelatinized hyphae, colored black
by the deposition of dark pigment-granules. The pseudopodetia are
covered by subglobose granules, showing the same color, morphology
endestructure as the granules of the thallus horizontalis (fig, 17) .
In some specimens a few stalks are branched.

fig. 8 P. ntgrtcaule (ce - cephalodium)

Pycnidia apical on short pseudopodetia or sessile on the thallus ho-
rizontalis. Conidiophores long, slightly branched with terminal
Sickle-shaped conidia, c. 10 x 1 pm.

Apothecia terminal on mature pseudopodetia or sometimes sessile
on the thallus horizontalis, 1.0-2.5 mm diameter. Apothecial margin
downturned as far as the point of attachment to the stalk. Apothe-
cia subglobose or slightly conical (fig. 8). No columella is present.
Apothecium and pseudopodetium separated by a broad boundary-texture.
No pigment-boundary is present. Hymenium c. 180 pm high, subhymenium
120 pm. Excipulum absent. Asci eight-spored. Spores rounded when
young, becoming spindle-shaped when mature, c. 18 x 7 pm.

Phycobiont green, Pleurococcus type, cells c. 10 pm diameter.

Cephalodia on the thallus horizontalis, thick, brown to black,

316

with wrinkled surface, c. O.5 mm diameter, containing Sttgonema.
Growing on Silicate rock.

Remarks

P. ntgrtcaule is distinguished from other species by the
white color and the thick, subglobose to peltate structure
of the thallus horizontalis and of the algal layer of the
pseudopodetium. With these characteristics the species is
easily dstinguishéd: from Pi veurtubims whieh as aiecorshor,
stalked and occurs in the same area. On the other hand, the
dark color of the central part of the pSeudopodetium, from
which the species takes its name, is not unique to the
species. The stalks of P. vegae, P. awasthtanum, P. curtulum, P.
strumatteus and P. eclavatus are also more or less blackened
within. As Kurokawa and Shibuicha (1970) pointed out, an
P. curtulum the color of the axis seems to vary with envi-
ronmental factors. The same applies to the other species
With blackened axes, where colorless and hyaline specimens
can always be found. The blackened axis appears to have no
consistent taxonomic value and therefore this criterion
should not be used to divide the genus Ptlophorus into sec-
tions as done by Sat6 (1940). The taxa Ptlophoron section
EupiLophoren "Sato (Journ. Jap.) Bou. 10: 175.040) eandeee—
lophoron section Wrorieaulia Sato. (hournmes Jap. boy 10%
ae 1940) therefore should be abolished.

Oe CESK

map. 9 P. ntgrteaule - distribution

oi 7

In figure 8 of his paper Sat6 (1940) gives a scheme of
the transverse section of pseudopodetium and apothecium
of his two sections. This rigure is misleading in several
respects. The species of his section Euptlophoron are shown
corbe hollow, Which 1s not true for most species of Prlo=
Pioris. Only 4ane Pr. aetcularte, and P. robustus do -old.pseudo-—
podetia sometimes become hollow. The section of Mgrtcaulta
shows a black central column of the pseudopodetium, which
passes directly into the central part of the apothecium.
In reality the apothecium and the pseudopodetium are not
epaly ot different ontogenetic origin, but are also dis-
tinctly separated by the boundary-texture. Furthermore,
the pigment is not restricted to the axis of the pseudo-
podetium, but the phyllocladia and the thallus horizonta-
lis are also pigmented.

Distribution (map 9)

The distribution of P. ntgrtcaule corresponds with the
occurrence of P. acteularte and P. clavatus, but the spe-
cies is much rarer. Until now P. ntgricaule has been found
in two areas only: Japan and the west coast of America.

Pilophorus robustus Th.Fr., De Stereocaults et Ptlophorts Comm.: 41
(1857). Type collection: Norway, Hedmark, Osterdal, Amot,
Austa-elv, 1837, Blytt (UPS, holotype).

Ptlophoron polycarpum Tuck., Am. Journ. Arts Sci. 2, 15: 427 (1858).
Type collection: Bering Straits (FH).

fig. 9 P. robustus (ce - cephalodium, p - pycnidium)

vy.

Thallus horizontalis granular or pulverulent, or indistinctly
squamulose, evanescent in older specimens, grey-green when dry. Gra-
nules about 1 mm diameter. Pseudopodetia stout, massive, seldom be-
coming hollow when old, irregularly or umbellately branched in the
upper part, normally 1.0-2.5 cm high, but sometimes up to 5 cm high
and 3 mm broad (fig. 9). Hyphae of stalk strongly gelatinized with
very small lumina (c. 0.5 pm), ecellishaty least. LO in length, ori-
entated more or less parallel to the longitudinal axis of the pseu-
dopodetium. Algal layer not homogenous, separated in loose granules.
Old pseudopodetia becoming more or less decorticate. Cortex-granules
c. O.2 mm diameter.

Pycnidia bottle-shaped, apical on the tips of short lateral bran-
ches or at the tip of smaller young pseudopodetia (fig. 9). Conidio-
phores c. 10 pm long, unbranched with elongate cells. Conidia ter-
minal, sickle-shaped, 5 x 1 pm.

Yoo rR

ete e ate e ee 2 ? A nN Both
map 10 P. robustus - world distribution (places listed @ and not
listed M® in the description. a specimens not examined)

319

Apothecia numerous, crowded at the apices of the pseudopodetial
branches, ustially’c. 2.5 mm (from 0.5 to 4 mm) broad, 1.5 mm high,
globose. Margins of crowded apothecia sometimes grown together se-
condarily. Margin of the hymenium extending downwards below the
point of attachment to the stalk, curving inward and upward before
reaching the pseudopodetial surface. Vegetative tissue forming a
large columella within the apothecium (fig. 12). In young specimens
only a small columella is present. Boundary-texture and pigment-
boundary absent. Hymenium c. 200 pm high, lower-half pigmented.

CHS 4
map Tl P. robustus - distribution in Scandinavia

320

Asci eight-spored. Spores simple, rounded when young, spindle-shaped
when mature, 18.0-24.0 x 4.0-6.5 pm. In one specimen some apothecia
without pseudopodetia sessile on the thallus horizontalis.
Phycobiont green, Pleurococcus type, cells c. 9 jam diameter.
Cephalodia large, brown to orange, lumpy, 1.0-1.5 mm diameter,
Situated on the thallus horizontalis and on lower parts of older
pseudopodetia (fig. 9), with Wostoc.
Growing on silicate rock and small stones.

Remarks

Ptlophorus robustus can be separated from all the other
species by the relatively tall, umbellately branched pseu-
dopodetia, the evanescent thallus horizontalis, and the
large and well-developed columella in the apothecium. In
some specimens the very young and small pseudopodetia re-
semble P. cereolus, but the latter is always sorediate. In
some cases certain specimens of P. robustus and P. acteu-
laris might be confused, especially in material collected
in Alaska, where both species occur together. This error
is possible if the pseudopodetia of P. acteularts are rich-
ly branched in their upper part and thereby resemble the
umbellately branched thalli of P. robustus. But the two
species can always be distinguished from the occurrence
of a columella in the apothecium of P. robustus.

Unusually small or very large specimens are quite fre-
quently encountered in this species. Some specimens from
different parts of the world are barely 1 ecm high and have
pseudopodetia of less than 1 mm in diameter. Others may
have pseudopodetia up to 5 em high and 6 mm broad at the
base. Among these*large specimens is P. robustus f. magnus
Savicz (1941),' but the type-material is in-no way unique |
among the specimens examined. It is very interesting that
unstalked apothecia which grew directly from the thallus

3 ® ie

SNe. Aker

Oy a s ee
b sry me 8 Sos, ee Os)
ovae SO PAL MESS
Pe SE Re Mig Se
SRT. a SS
ZA =| a Soe ‘ge be ——

—— = aS Imm
fig. 10 P. strumatteus (ap - primordium of apothecium,
ce — cephalodium)

horizontalis were found. This is quite common in species
with small pseudopodetia such as P. ftbula, but is rather
unusual in the larger species.

Distribution (map 10-11)

WAAL

Pe ropustus «snows -ardastinct arctic, circumpolar distri-

bution. In Scandinavia where most samples were found the

Species is almost exclusively restricted to the high moun-

tains as was already established by Ammann and Ammann
(1969); but here too, its clear dominance in northern re-

Plonseis, Obvious. Alsecond centeryor distribution aseAlas—

ka and the Tschuktschen Peninsula. These two main areas
are connected by specimens collected in Siberia, Spits-
bergen and Greenland. I have not seen the material from
Spitsbergen myself, but the information was given by Krog
Specimens found outside Scandinavia and Alaska:
Spitsbergen, Svenskegya, Kong Karlsland, E.Dahl, 1936; Greenland,
Laksefjord (C); Greenland, Laksefjord, Proven District (C); Green-
land, Diskofjord, Kuanersuit (C); Greenland, Disko, Nordfjord,
Kugssinerssuaq (C, 2 specimens); USA, Alaska, Aleutian Islands,
Amchitka Island (WIS); USSR, Tschuktschen Peninsula (2 specimens
LE, M); USSR, Arakamchechen Is., Tschuktschen Peninsula (BM, FH, PC,
US); USSR, Wrangel Island (3 specimens LE); USSR, Taimyr Peninsula
(3 specimens LE; GB, 2 specimens S); USSR, Laja River, Bolchese-
melskaja (LE); USSR, Tobolsk (LE); USSR, Beresow, near Tobolsk (LE,
Oa

Pilophorus strumaticus Ny]. ex Cromb., Monogr. Lich. Brit.: 115
(1894). Type collection: Great Britain, S. Aberdeenshire, Brae-
mar, Morrone, 1869, Crombie (BM, lectotype).

P. robustus f£. dtstans Hult., Bih. K. svenska Vetensk. Akad. Handl.
26: 17 (1900). Type collection: Sweden, Dalsland, Hattefjdll,
prope Vagsdter, 1870, Hulting (GB, lectotype; UPS, lectopara-
type (1895)).

P. dtstans (Hult.) Magn., Bot. Notiser 108: 298 (1955).

Thallus persistent, minutely granular or nearly squamulous; gra-
nules O.1 mm diameter, often becoming confluent to form aggregations
c. 1 mm in size, grey when dry. Pseudopodetia pin-like, up to 2 mm
high, O.5 mm diameter (fig. 10). Internal hyphae strongly gelati-
nized, cell lumina O.5 wide, hyphae of upper part usually grey-
brown. Pseudopodetia mostly covered by a layer of algae in con-
tinuous granules or squamules.

-Pycnidia apical on short pseudopodetia, up to O.5 mm high, often
surrounded by a collar of granules or cortical scales. Conidiopho-
res c. 45 pm long, unbranched with terminal sickle-shaped conidia
Dex il pmein’ size.

Each mature pseudopodetium nearly always bears a single hemi-
spherical apothecium, c. 1 mm diameter. In some cases two separate
apothecia develop at the tip of the pseudopodetium but later merge
together. Apothecial margin down turned as far as the point of
attachment to the stalk. The boundary between the generative and
vegetative tissue arches upwards to form a small columella at the
center of the apothecium (fig. 12). The boundary-texture and pig-
ment-boundary between the generative and vegetative tissue are ab-
sent in mature apothecia. The development of apothecial primordia,
which have a pigment-boundary is very characteristic for the spe-

Sa7

cies (Jahns, 1970a).

Hymenium c. 230 pam high, lower part pigmented, margin sterile,
consisting only of paraphyses. Excipulum absent. Asci eight-spored.
Spores rounded when young, becoming spindle-shaped when mature,
TATOS19, 08X46. SS 25 pm. The brown-black pigment is densest directly
below the paraphyses and tends to become less intense towards the
columella.

Phycobiont green, Pleurococeus type, cells c. 8 pm diameter.

Cephalodia brown, granular, surface irregular and lumpy, contain—
ing Sttgonema. The development of the cephalodia was described else-
where (Jahns, 1972).

Growing on silicate stone.

map 12 P. strwmaticus - distribution in Great Britain

EAE:

Remarks
The distinct columella in the apothecium is character-

bstic Of Mature specimens of Chis species,. distinguishing
them from the other small species. P. robustus the only
other species with a columella, has a distinctly different

habit.

map 13 P. strumatteus - distribution in Scandinavia ( mm places
not listed in the description, A specimens not examined)

324

The name P. strumattcus was first used by Crombie (1875,
p. 140), but as no description was appended it was a nomen
Udine Cine re.

Distribution (nap 12-13)

P. strumatitcus has been found in western Scandinavia and
in Great Britain. In contrast to P. robustus and P. cereolus,
this species obviously prefers an oceanic climate and ex-
tends far less to the east and north of Scandinavia than
the other two species. Similar observations were made by
Ammann “and, Ammann (19090)\eeTwo of their locations have
been included in map 13 (marked. by a triangle), without
having been examined by me.

Pilophorus vegae Krog, Norsk Polarinstitutt Skrifter, 144: 56-57,
Oslo (1968), fig. 14. Type collection: Alaska, Bering Sea
district ,sNunivak ishand,. Roberts Mt. vl7O0.ft, Kroqt(0;,
holotype).

Thallus horizontalis crustaceous, consisting of loosely aggre-
gated or nearly evanescent granules, c. O.5 mm diameter, white when
dry. Pseudopodetia simple or branched in the upper part. Branches
either dichotomous or irregularly bush-like. Pseudopodetia 1.0-1.5.
cm high and c. 3 mm broad. Younger parts covered with a continuous
algal layer, consisting of minute granules or scales. Color of
scales white, later becoming blackened at the base of the pseudopo-
detia. Basal part of old pseudopodetia often ecorticate. Central
column of the young stalks hyaline, while the interior of older
stalks is completely black.

fig. 11 P. vegae (ap - primordium of apothecium, ce - cephalodium,
s - broken stalk with pigmented center)

RES

Pycnidia not observed.

Mature apothecia are absent in all specimens examined, but pri-
mordia of apothecia are found terminal at the tip of the stalks.

Phycobiont green, Pleurococcus type.

Cephalodia stipitate on the pseudopodetia (fig. 11), containing
Nostoe in the type-specimen and Sttgonema in other specimens. Small
cephalodia-like structures on the substrate contained algae of the
Gloeocapsa type, but it seems doubtful, whether they belong to
this lichen.

Growing on silicate rock.

Remarks
P. vegae can be confused with P. robustus and P. actcula-
rts occurring in the same region. The clearest character-
istics of the species, which are lacking in P. robustus
and P. acteularts, are the white color of the young pseudo-
podetia and the black pigment of the central column. P.
ntgrtcaule, which also has a white cortex and a black cen-
tral column is smaller and easily recognized by the glob-
ular or peltate shape of its cortex granules.

map 14 P. vegae - distribution ( e@ places listed in the des-
cription of the species)

52:0

Distribution (map 14)

P. vegae is known from Alaska and the Bering Strait
area only. I have seen the following specimens:
USSR, Tschuktschorum, Bovidential, 64°45'N, 174°w (2 specimens LE,
O); USA, Alaska, Bering Sea district, Nunivak Island, Roberts Mt.,
1700 £t(O); USA, Alaska, Central Pacific Coast, Talkeetna Mts. (0);
Canada, British Columbia, Queen Charlotte Islands, Moresby Island,
Laing Point Mountain, SE of road into Peel inlet, c. i mile of Laing
Renliae,; LOO) ae (GAN) ~

Lichens to be excluded from the genus Pilophorus

Four species have been described as belonging to the
genus Ptlophorus, which must be placed elsewhere:

Pilophorus colensoi (Bab.) Knight, Trans. Proc. New Zealand 16: 400
(1884).
Stereocauton colensot Bab., in Hooker, Flora N.Z. 2: 294 (1867).
Type collection: New Zealand, Colenso 2746 (BM, holotype).
Corynophoron colensot (Bab.) Nyl., Lich. Nov. Zeland.: 15 (1888).
Thallus horizontalis thin, granular and evanescent. Pseudopode-
tia up to 5 mm high, branched. Cortex of irregular scales. Cepha-
lodia sacculate, on the pseudopodetia, containing Sttgonema. De-
velopment of apothecia following the typical ontogeny of a Stere-
ocaulon species (Jahns, 1970a).

Remarks

The helical ascogones are especially typical and quite
different from the straight ones found in the genus Prlo-
phorus. The species therefore belongs to the genus Stereo-
caulons @ View salso taken by Th.Fries (les7) and aby mam,
Lamb in a note attached to the type-specimen.

Raésdnen (1932, p. 24) described P. colensot var.reagens
Ras. from Tierra del Fuego.

I have seen material from the following places:
New Zealand, Colenso 2746 (BM, holotype); Fuegia occid. supra Ru-
pium (B); N-East of New Zealand (G).

Pilophorus conglomeratum F.Wils., Linn. Soc. Journ. Bot. 28: 372
(1891). Type collection: Hab. supra truncum arboris smortuum
muscosum in Black spur. Victoria, Australia (BM, holotype).

Ptlophorus cartosum Hue, Nouv. Arch. Mus. 3, 10: 280 (1898).

Thallus horizgontalis thin, grey-green, consisting of minute gra-
nules up to O.1 mm in diameter. Thallus horizontalis persistent or
evanescent. Thallus verticalis up to 2 cm high, pin-like, un-

branched. In young specimens more or less smooth, covered with a

thin layer of minute granules similar to those forming the thallus

horizontalis. Thallus verticalis quickly becoming bare and divided
by lengthwise-oriented furrows. In old specimens the stalk consists
of numerous nearly-separated columns.

fig. 12-16 anatomy of apothecia (bt - boundary-texture, c - colu-
mella, pb - pigment-boundary)

Tig. P. ntgrtcaule with peltate scales, longitudinal
section of the stalk

Sat
LED
Sey
ny
ay
ay
ba
Sw
Say
a
Seng
~
=;
fy
S
—

=
=
Ss
=
=
—
——
Ss
—
Fs
—
ZS
¢
Ss
S
“a

i( 3
Noy
16 1%,

DLO

Several small apothecia crowded at the tip of the stalks, gradual-
ly merge into one large fruitbody. Spores 14 x 3.5 pm, spindle-
shaped. Apothecia dark-brown. Cephalodia absent.

Remarks

The species somewhat resembles P. robustus, but the ab-
sence of cephalodia and the brown pigment of the apothe-
cia are striking differences. The whole habit is slightly
different from a Ptlophorus and therefore the species
should be excluded from the genus. Perhaps the lichen is
related to Baeomyces or to one of the genera of the Clado-
niaceae (e.g. Heteromyces, Thysanothectum) but the question can
not be settled before more is known of its ontogeny.
I have seen material from the following places:
Tasmania, Hartz Mountain plateau, on rotten wood in forest, extreme-
ly damp and well shaded, alt. 3700 ft (BM); Central Western Tasma-
nia, forest at western side of Mt. Arrowsmith near Franklin River,
(BM); North Western Tasmania, Ball Room Forest Track, Dove Lake,
near Cradle Mt. (BM); New Zealand, South Island, Fjordland, Eglinton
Valley, Cascade Creek, alt. 1600 ft (BM); Victoria, Australia (BM,
holotype) .

Pilophorus staufferi Frey (nomen nudum), Bot. Jahrb. 86, 1-4: 242
(1967). Type collection: Neukaledonien, Monts des Koghis, Mt.
Bono, Gipfelgrat, 1070 m (BERN).

One's first impression of this species is that 16 as
very much like P. robustus. The thallus verticalis is um-
bellately branched in the upper part, each branch bearing
one to several apothecia. But on close inspection the
lichen seems to be related to P. conglomeratum. The color
of the apothecium is more brown than black, and the stalks
Show the same furrows as in P. conglomeratum. Cephalodia are
also-absent. Thelthallus horizontalis of —Peterausrerr, .un=
like the granulated thallus of P. conglomeratum, consists
of small phylocladia about 1 mm in diameter. I think that
P. stauffert and P. conglomeratum belong to the same genus;
their exact taxonomic positions must be decided later.

Pilophorus pileatum (Mont.) Zahlbr. (nomen nudum), Catal. Lich.
1023S) Se (1939)e

Cladonta ptleata Mont., Annal. Scienc. Nat. Bot. 3, 18: 310 (1852)
Type collection: Coquimbo in Chile.

I have seen no samples from this species, but the
descriptions of Montagne (1852) and Vainio (1887) mention
brown-black apothecia. Therefore it certainly does not
belong to Ptlophorus.

References

Acharius, E. (1798) Lichenographiae Sueciae Prodromus. Lincopiae.
Acharius, E. (1803) Methodus qua omnes detectos Lichenes. Stockholm.
Acharius, E. (1810) Lichenographia universalis. Gdédttingen.

Ammann, K. & B. Ammann (1969) Die fennoskandische Verbreitung von
Pilophorus (Tuck.) Th.Fr., Stereocaulaceae. Herzogia 1:87-94.
Crombie, J.M. (1875) Recent Additions to the British Lichen-flora.

J.Bot.Lond. 13:140-142.

SENS)

Crombie, J.M. (1894) A Monograph of Lichens found in Britain.
British Museum, London.

Culberson, W.L. (1970) The Typification of the Lichen Genus Pilo-
phoron. Bryologist 73:630-632.

Fink, B. (1910) The Lichens of Minnesota. Contr. U.S. nat. Herb.
14, 1:1-269.

Frey, E. (1967) Die lichenologischen Ergebnisse der Forschungsreisen
des Dr. Hans Ulrich Stauffer. Bot. Jahrb: 86, f-4:3209-—255.

Fries, Th. (1857) De Stereocaulis et Pilophoris Commentatio. Uppsala.

Fries, Th. (1861) Lichenes Arctoi Europaeae Grénlandiaeque hactenus
COgniti.. Nova, Acta Reg ..soc. scient.,Uppsal.. 3,°322/8;

Fries, Th. (1871) Lichenographia Scandinavica I, Uppsala.

Fries, Th. (1874) Lichenographia Scandinavica II, Uppsala.

Fries, Th. (1888) Nagra anmarkningar om Sladgtet Pilophorus.

Bot. Notiser 1888: 212-214.

Fries, Th. (1889) Einige Bemerkungen Uber die Gattung Pilophorus.
Bot. Centralblatt 38:764-766.

Hawksworth, D.L., P.W. James & J.R. Laundon (1972) The Nomenclature
OrePilopnorusc., Taxon 21 ,52/ 5532/2920.

Henssen, A. & H.M. Jahns (1973) Lichenes. Stuttgart.

Hooker, J. (1867) Handbook of the New Zealand Flora II. London.

Hue, A.M. (1898) Lichenes extra-europaei. Nouv. Arch. du Muséum 3,
10:280

Hulting, J. (1900) Dalslands Lafvar. Bih. K. svenska Vetensk. Akad.
Handl.1.626,00/3:17.

Jahns, H.M. (1970a) Untersuchungen zur Entwicklungsgeschichte der
Cladoniaceen unter besonderer Berticksichtigung des Podetium-
Problems. Nova Hedwigia 20:1-179.

Jahns, H.M. (1970b) Remarks on the taxonomy of the European and
North American Species of Pilophorus Th.Fr. Lichenologist 4:199-
213.

Jahns, H.M. (1972) Die Entwicklung von Flechten-Cephalodien aus Sti-
gonema-Algen. Ber. Dtsch. Bot. Ges. 85, 10-12:615-622.

James, P.W. (1965) A new Check-list of British Lichens. Lichenolo-
GSC ess Eo=i 53.

Knight, C. (1884) On the Lichenographia of New Zealand. Trans Proc.
New Zealand 16:400.

Krog, H. (1968) The Macrolichens of Alaska. Norsk Polarinst.
Skrifter 144:1-180.

Kurokawa, S. & H. Shibuichi (1970) Notes on Japanese Species of
Pa OpnOron. WOurn. sap. Sot. 45, 2 /5=<04.

Magnusson, H. (1937) Férteckning 6ver Scandinaviens vaxter, 4 Lavar.

Magnusson, H. (1955) New or otherwise interesting Swedish Lichens
XV. Bot. Notiser 108:298-299.

Montagne, D.M. (1852) Diagnoses phycologicae. Ann. Sci. Nat. 3, 18:
302-319.

Nylander, W. (1857) Enumération générale des lichens. Mém. Soc.
Sci. nat. Cherbourg 5:85-146.

Nylander, W. (1860) Synopsis Methodica Lichenum I. Paris.

Nylander, W. (1888) Lichenes Novae Zelandiae, Paris.

Rasanen, V. (1932) Zur Kenntnis der Flechtenflora Feuerlands. Ann.
Socs Zool. Bot. Fenn. Vanamo 2, 1:24.

Rasdanen? V. (1946) Lichenes novi II. Ann. Soc. Zool. Bot. Fenn.
Vanamo,, Notil. ©Bot. 2isi=7.

Rasanen, V. (1950) Lichenes novi VI. Ann. Soc. Zool. Bot. Fenn.
Vanamo 5:28-29.

330

Sat6, M.M. (1933) Notes on some Japanese Lichens determined by Dr.
Bav. A.) Wainio Lil. Journ. Jap. Bot. 9221i-214.

Sato, M.M. (1940) East; Asiatic) Lichens ITi< \ Journ. Jap. Bot. 16:
173-147:

Savicz, Vee. (1941) De nonnullis lichenibus URSS. Notul. syst. inst.
GLY DECG .sHOrLILSDOLtEreLLOpol. =o 4/—9".

Stitzenberger, E. (1862) Beitrag zur Flechtensystematik. Ber. Utiber
die Tatigkeit St. Gallisch. naturwis. Gesellsch.:124.

Tuckerman, E. (1847) A synopsis of the Lichens of the Northern Uni-
ted States and British America. Proc. Am. Acad. Arts Sci. 1:

195-285.

Tuckerman, E. (1858) Supplement to an Enumeration of North American
Aken, Nm dobien, Braces Serle, Be Bos422-a3©0,

Tuckerman, E. (1877) Observationes Lichenologicae 4. Proc. Am. Acad.

ADESSO a2 is ie

Tuckerman, E. (1882) A Synopsis of North American Lichens. Boston.

Vainio, E.A. (1887) Monographia Cladoniarum universalis.

Vainio, E.A. (1921) Lichenes ab A. Yasuda in Japonica collecti,
Continuatio 1. Bot. Mag. Tokyo. 35:45-79..

Wilson, F..(1891) On Lichens collected in’ the Colony of Victoria,
Australia, Linn.) Soc, Journ, Bot. 2e-3i2.

Zahlbruckner, A. (1926) Lichenes. In: Die nattirlichen Pflanzenfa-

milien, 72., Autl., Bd.S, nrsGg. von englen, A. S)Kk.) Prantls.beipzig.

Zahlbruckner, A. (1927a) Additamenta ad Lichenographiam Japoniae.
Bot. Mag. Tokyo 41:313-364.

Zahlbruckner, A. (1927b) Catalogus Lichenum Universalis 4. Leipzig.

Zahlbruckner, A. (1939) Catalogus Lichenum Universalis 10. Leipzig.

MYCOTAXON

Otel NO) DD. wos 558 July-September 1981

NEW DEMATIACEOUS HYPHOMYCETES
FROM) TROPICAL, RAIN FOREST LITTER

S. ONOFRI, D. LUNGHINI, A. RAMBELLI and L. LUSTRATI

Cattedra di Micologia, Istituto dell'Orto Botanico, Universita
di Roma, Largo Cristina di Svezia 24, 00165 Roma, Italy.

Summary

In this paper, anew genus, Chryseidea gen.nov. with the new species
C.africana, and two new species, Beltrania onirica sp.nov. and Phaeo-
trichoconis aurata sp.nov., from Tai National Park (Ivory Coast) fo-
rest litter, are described.

Among mycological investigations carried out by this Chair on the
Tai project (M.A.B., Project n.1) to study the influence of human ac-
tivities on the south-west forests in the Ivory Coast and with the pur-
pose to improve the environmental knowledge and quality, three new
species and a new genus of Dematiaceous Hyphomycetes are described.
They were found on dead leaves collected on the soil surface in the Tai
National Park forest. Every attempt at isolation in pure culture was
unsuccessful. The following descriptions concern observations on na-
tural media,

Exsiccata are available in the Mycological herbarium of the Rome
Botanical Garden, (RO).

Chryseidea Onofri gen.nov.

Conidiophora macronematosa, synnematosa, erecta, simplicia, pig-
mentata, in apice semper sterilia, Cellulae conidiogenae a latere po-
sitae, determinatae, discretae, simplices, ampulliformes, apice den-
ticulato, circiter in medio synnemati circumpositae, Conidia hyalina,
falcata,

Sp.typ.: C.africana Onofri sp.nov.

Conidiophores macronematous, synnematous, straight, simple, pig
mented, with apices always sterile. Conidiogenous cells lateral, de-
terminate, discrete, simple, ampulliform, with a denticulate apex,
surrounding the conidiogenous apparatus approximately in the mid part.
Conidia hyaline and falcate,

40 um i

rs
——
—Y
==

=——

ee
ee

——

SSS

Nk
h

555

Chryseidea africana Onofri sp.nov.

Conidiophora macronematosa, synnematosa, solitaria, recta, simpli-
cia, adtenuato et sterili apice, ex constantibus et rectis hyphis (2 um
crassis) composita, pallide brunnea, usque ad 560 wm longa et 42 um
ad basim crassa, Cellulae conidiogenae a latere positae, discretae,
simplices, ampulliformes, cum apice saepe deflectens conidiorum
forsitan pondere vel magna copia et denticulato ob subsequentes loco-
rum qui conidia generant proliferationes; orientes ex hyphis laterali-
bus, adherentibus et parallelis ab hyphis synnematis et orientibus pro
xime sub saeptis; ab circiter tertio altitudinis a basi usque ad dimidium
synnema circumpositae, sufflavae; 7,8-11,2x2,3-2,7 wm. Conidia fal-
cata, asaeptata, extremitatibus acuta, hyalina, omnia agglutinata; cir
cum conidiophora permanent in formam similia Chaetopsina fulva;
14,5-18,5x1,4-1,9 wm.

In foliis emortuis, Tai, Ora Eboris, holotypus: H.B.R. 116A, (RO).

Conidiophores macronematous, synnematous, solitary, straight,
simple, narrowing to the sterile apex, composed of straight and regu-
lar hyphae, Ochraceous-Tawny in colour (Ridgway, 1912), up to 560
um long and 42 wm thick at the base, Conidiogenous cells lateral,
discrete, simple, ampulliform and denticulate from successive proli-
ferations, with the apex frequently repent, presumably under the weight
of conidial production; they originate from lateral hyphae that grow
just under the septa of conidiophorous apparatus at about one third of
the height from the base and extend up to half of the height of the coni-
diophorous apparatus, along which they grow side by side, Light-Buff
in colour (Ridgway, 1912); 7,8-11,2x2,3-2,7 wm. Conidia 0-septate,
falcate, with very sharp apices, hyaline, 14,5-18,5x1,4-1,9 wm; they
remain coiled at the apex of each conidiogenous cell in a compact mass
surrounding the conidiophorous apparatus.

The microorganism described presents morphological characters
different from others belonging to genera with synnematous conidio-
phorous apparatus, It differs in the position of the conidiogenous cells
along the conidiophorous apparatus, in the resulting general structure
(Chaetopsina like) and in spore shape. For these considerations we
think necessary to propose the new genus, Chryseidea, and the spe-
cies C,africana for our strain,

Beltrania onirica Lunghini sp.nov.

Coloniae effusae, Mycelium immersum, Setae erectae, simplices, le-
ves, atrobrunneae, usque ad 506 um longae et 4,6 wm ad basim crassae,
Conidiophora macronematosa, mononematosa, simplicia aut rarius ra
mosa, saeptata, flexuosa, levia, non constanti crassitudine, pallide
brunnea, 184-236x4,6 um; ab eadem basi setarum oriuntur, Cellulae
conidiogenae integratae, terminales, polyblasticae, denticulatae, sym

334

20 wm

40 um

a. conidiophores; b, conidiophores

Fig. 2 - Beltrania onirica sp.nov.:

and seta; c. conidia.

podiales, locis qui conidia generant magno intervallo positis, pallide
brunneae. Conidia levia, acrogena, protoplasmate granuloso, solitaria,
conica-campanulata, zona supraaequatoriali subhyalina et tenui orna-
ta, ad basim denticulata, ad apicem breviter appendiculata, pallide
brunnea; 19-21x11,5-12,5 wm; appendix circiter 2 wm longa.

in foliis emortuis, Tai, Ora Eboris, holotypus: H.B.R..117A, (RO).

Colonies effuse. Mycelium immersed. Setae straight, simple, with
thick and smooth walls; Argus Brown in colour (Ridgway, 1912); length
up to 506 wm and 4,6 wm thick, Conidiophores macronematous, mono-
nematous, simple or rarely branched, septate, slightly curved and
irregulary thick, Deep Olive-Buff in colour (Ridgway, 1912), 184-236
x4,6 wm; originating from the base of the seta. Conidiogenous cells
integrated, terminal, polyblastic, denticulate, elongating by sympo-
dial proliferation, Deep Olive-Buff in colour (Ridgway, 1912).

Conidia smooth, acrogenous, with granular protoplasm, solitary, co--
nic-campanulate, with a thin, not easily observed transverse band,
with a very short appendage at the apex and denticulate at the base,
Deep Olive-Buff in colour (Ridgway, 1912); 19-21x11,5-12,5 wm; appen
dage about 2 wm long.

The microorganism described presents characters different from
those of related species, In particular it differs from Beltrania querna
Harkness (Pirozynski, 1963) with conidia with a different shape and
longer appendage; the conidial shape of our strain is very close to
those of B, mangiferae Munjal et Kapoor (Pirozynski and Patil, 1970)
and B, muelleri Rao et Mani Varghese, 1978, but these species are
characterized by different conidiogenous cells, For these considera-
tions we propose the new species B.onirica for our strain.

Phaeotrichoconis aurata Rambelli sp.nov.

Coloniae effusae, Mycelium immersum, Conidiophora macronematosa,
mononematosa, solitaria, erecta, simplicia, saeptata, levia, brunnea-
sufflava, usque ad 126 wm longa et 3,6-5 wm crassa (ad basim 8-10 wm),
Cellulae conidiogenae polytreticae, integratae, sympodiales, cicatri-
cibus conidialibus parum manifestis preditae, Conidia solitaria, sicca,
acrogena, obclavata, 5-saeptata, ad basim trunca, ad apicem longo
rostro praedita, levia, pallide brunnea-flava, protoplasmate omogeneo;
34,5-43,7x4,6-6,9 um; rostrum rectum, asaeptatum, hyalinum, usque
ad 25 um longum et usque ad 2 wm crassum,

In foliis emortuis, Tai, Ora Eboris, holotypus H.B.R. 118A, (RO).

Colonies effuse, Mycelium immersed, Conidiophores macronematous,
mononematous, solitary, straight, simple, septate, with smooth walls,
Buffy Brown in colour (Ridgway, 1912), up to 126 wm long and 3,6-5 wm
thick (8-10 um at the base), Conidiogenous cells polytretic, integrated,
sympodial, Conidia solitary, dry, acrogenous, obclavate, transversely

———_—— Oe

356

Fig. 3 - Phaeotrichoconis aurata sp.nov.: conidiophores with conidia,

Koy ii

SRO acc

Sette rte =

sco saieneatensnnpeeceremnenraenstitene rgesesian

Fig. 4 - Chryseidea africana: a. and b.
Phaeotrichoconis aurata: c. and d.
Beltrania onirica: e, and f,

338

5-septate, rostrate, with a large scar at the base, smooth, Tilleul-
Buff in colour (Ridgway, 1912), with homogeneous protoplasm, 34,5-
43,7x4,6-6,9 um; scar about 2,5-3 wm wide, beak straight, aseptate,
hyaline, up to 25 wm long and 2 um thick,

Ellis (1971) modified the original diagnosis of the type species
P, crotalariae (Salam et Rao) Subramanian to include also species
characterized by polytretic, integrated, terminal and sympodial co-
nidiogenous cells, Our microorganism seems includable in the genus
Phaeotrichoconis as modified by Ellis (1971), but differs from the two
described species, P,crotalariae and P,urariae Bharadway, in seve-
ral characters and mainly in the dimensions; for these reasons we
propose for our strain the name P, aurata as a new species.

ACKNOWLEDGEMENTS

The Authors wish to thank the Minister of Scientific Research of
the Ivory Coast, for the assistance he has given, Dr. P.M. Kirk of
the C.M.I. for a slide of P.crotalariae kindly sent us and our collea-
gues, in particular Dr. A.M. Persiani, of the Laboratory of Mycology
for their criticisms. We also thank Prof. O. Verona, Istituto di Micro
biologia Agraria eTecnica, Universita di Pisa, Italy, for kindly revie-
wing the manuscript.

REFERENCES

Bharadway, S.D. (1969). A new species of Phaeotrichoconis on Ura-
ria picta from Varanasi, India. Indian Forester 95(1): 24-26.
Carmichael?) [,W:, W.B.° Kendrick, “Isl. Connerswande py. o1glerg( 1980).

Genera of Hyphomycetes. The University of Alberta Press, Edmon-
ton, Alberta, Canada.

Ellis, M.B. (1971). Dematiaceous Hyphomycetes. Commonw. Mycolog.
Inst., Kew, England.

Ellis, M.B. (1976). More Dematiaceous Hyphomycetes. Commonw. My-
colog.=Inst., Kew, England.

Matsusnhimaset = .(L0/))- Microfungi of the Solomon Islands and Pa-
pua-New Guinea. Publ. by the Author, Kobe, Japan.

Matsushima, T. (1975). Icones microfungorum a Matsushima lectorum.
Publis byethie, Author mKobDewaapan.

Morris, E.F. (1963). The synnematous genera of the Fungi Imperfec-
ti. Western sillinois| Univ; {Ser sbiclogicalmociencesmio ws).

Pirozynski, K.A. (1963). Beltrania and related genera. Mycological
Papers, 90.

Pirozynski, K.A. and S.D. Patil (1970). Some setose Hyphomycetes
of leaf litter in south India. Can. J. Bot. 48: 567-581.

Rao, V.G. and K.I. Mani Varghese (1978). Litter-Fungi: two unde-
scribed Hyphomycetes from India. Nat. Acad. Sci. Letters 1(2):
49-50.

Ridgway, R. (1912). Color standards and color nomenclature. Wash-
ington, 1b.CG.

Subramanian, C.V. (1971). Hyphomycetes, An account of Indian spe-—
cies, except Cercosporae. Indian Council of Agricultural Research,
New Delhi, India.

MYCOTAXON

VORPENTDIC RENO S825 epp5ji559=545 July-September 1981

COCHLIOBOLUS RAVENELIT SP. NOV. AND C. TRIPOGONIS SP. NOV.

J. L. ALCORN

Department of Primary Industries,

Indooroopilly, Queensland, 4068, Australta

Abstract

Cochltobolus raveneltt sp. nov. and C.
trtpogonts sp. nov. are described from culture,
as the teleomorphs for Btpolarts ravenelit and
B. tritpogonts comb. nov. respectively.

Pompe Vimo (ames LUC Vaso EEpenenCsCOnCceptsein ticacomplex
of fungi assigned to Drechslera sens. lat., I have attempted
to produce teleomorphs for species where no such connection
was known. This has been done to test the hypothesis that
conidial state characters allow prediction of teleomorph
correlations, and that new associations will prove to be
Drechslera sens. str. - Pyrenophora, Btpolarts - Cochltobolus,
and Exserohtlum - Setosphaerta (Luttrell 1977, 1978). Three
new teleomorphs were described previously, two in Cochlto-
bolus (Alcorn 1978a) and one in Setosphaerta (Alcorn 1978b),
and those results support the predictive aspect of anamorph
classification. Subsequently Cochltobolus states have been
induced to form in cultures of other species. Two of these
teleomorphs are described here as new.

COCHLIOBOLUS RAVENELIIL

Ascal states in Cochltobolus were found by McKenzie
(1968) for two 'Helminthosporium' species which infect
Sporobolus inflorescences in Queensland. The work was
reported in a thesis, and subsequently none of the data
presented have been published. McKenzie did not assign
specific epithets to the Helminthosporium collections

340

studied. I have examined specimens she deposited in BRIU
and there is no doubt that one of the anamorphs connected
with a teleomorph was Bipolarts ravenelit (Curt.) Shoem.
Luttrel1 (19:76) was unable to produce a-per tech State rior
this species, and I made many unsuccessful pairings of
cultures before a teleomorph developed. The methods used
in the successful attempt are described briefly below.

Ten collections of Sporobolus spp. with infections of
the inflorescence caused by B. ravenelit were made in south-
eastern Queensland, in an area approximately 60 km north of
Brisbane. Conidia from each of five panicles in each
collection were streaked over water agar plates and allowed
to verminate., Pour verminatecasconidiamtrom cachap aveswere
transferred to PDA, making a total of 20 transfers for’ each
coblection. “Cullturesaweresparred=on sachs agar (hevere
1971).) each plate being amended with» maize lear sand barley
Uraim. sihrcenmethodss0t. pairing. weresecmployed,

A. Bulked conidial inoculum was prepared for each
collection. Conidial suspensions from all isolates
from a single collection were mixed and inoculated
onto the plates.

B. All possible pairings: bDetweenscollectwons weresmade,
using the bulked conidial inoculum from each.

C.. “All possable pairings between collections, using
one single-conidium isolate from each. Inoculum
consisted of blocks from PDA cultures.

Ascocarps were obtained in only one pairing, namely
with bulked inoculum from collection 7957 (Sporobolus
ferttlts (Steudel) W.D. Clayton, Glasshouse Mountains,
24.v.1979) and collection 7963 (S. elongatus R. Br., .Maleny.,
J4.V,19/9)4, “Pour ascocarpsuwere: formed, each iwi tit aso LOpose
basal’ body and a long cylindrical neck. They were visible
only because the necks projected above the dense hyphal mat
investing the basal section. Ascocarps contained many ripe
asci which released ascospores freely in water. Single-
ascospore culturés were established. These cultures were
paired in all possible combinations, but only one pairing
resulted in the formation of ascocarps. None were formed
in single-ascospore cultures, but all produced an anamorph
on PDA indistinguishable from B. ravenelit.

34]

In another test, small numbers of ascomata formed in
Dalringspotucultures. fromecollections 79/6 (5.7 fertiiis,
St. Lawrence, 24.vii.1979) and 7977 (Sporobolus sp.,
Claliviewm 4 .Vi119 79). andy /976.and 79/89(S. fertilie,
between Sarina-and Mackay, 24.vii.1979). Some single-
ascospore isolates from these ascocarps produced the teleo-
morph when paired, but not when grown alone under the same
cultural conditions. Subsequently ascomata were formed in
greater numbers when inflorescences and grain of S. elongatus
Were Used iasathe plant substrate ian oach"s agar plates,
mnsteadeotemarze, lean or Darley agradin:.

Cochltobolus ravenelit sp. nov. Figures 1 - 4

Ascocarpi atrobrunnei vel atri, basi globosa 360-515 um
diam. et collo cylindrico 980-1800 um alto, ad basim 95-135
DMparall eae CON so ouiine G1 alemAS Cia cy WIndri Ca.
Vestigialessbitunicatdi , 165-500)x 14-1/-5 im... /AScosporae
hyalinae, filiformes, versus extrema angustatae, rectae vel
torsivae, 5-14-septatae, 155-310 x 5-7.5 um.

In inflorescentiis et granis Sporobolt elongatt in
AC ATOM d Chile ile OSU] BRI Pals LOomenolOLY DUS em DRUP 5027),
IS0ZSeparatypi .

Ascocarps dark brown to black with a globose body 360-
J Omid lalimeanded paler cylindrical neck 980=1800) x 95-
IS oeiMeate chesbase and) 95-1 loimeat. thesapex = Occasionally
with a cylindrical base up to 280 um x 250 um below the
ascogenous section. Ascocarp body often covered densely by
reddish brown hyphae, the beak glabrous or with a few short
Nag TSeiGl oie SASCIMCYINGtICal wUSidallyetanered «si only
near the base but. lacking a distinct, pedicel, straight vor
eurveds 9165-4002 x 114-175 ime Often, with asblunt baturcate
foots o- Om median... VeSit1 ol alebitunicate Ur lens 2)/0 mASCOs
Sporesuri ii tom: Stralont) ovecoi led helvcaluiys in thepascus.
slightly tapered to obtuse apex, more so to base, 5-14-
Ceptate loo LUE Xo oe Time Ore)

| In some asci the degree of ascospore coiling is only
very slight, while in others it is more pronounced. Some-
times ascospores are tightly coiled in the apical part for
20-25 um, and loosely in the remainder. Ascospore diameter
at the apex is about 4 ym, and at the base 2.5 wm. Conidia
from two collections which gave rise to inter-ferile
cultures of B. raveneltt are illustrated in Figure 4.

342

Cochltobolus ravenelii is quite distinct from C.
sporobolt Castellani, which occurs on Sporobolus leaves
(Castellani 1951). C. sporobolt has smaller, short-beaked
ascomata, smaller asci, and shorter 4-6-septate ascospores.

COCHLIOBOLUS TRIPOGONIS

Drechslera trtpogonts A.S. Patil & V.G. Rao was
described from India, infecting the inflorescences of
Tritpogon jacquemonttt Stapf (Patil §& Rao 1972). Subsequently
L-collected a similar fungus onl. cOLt7) Ormes etl eMuel 1}
C.E. Hubbard in Australia (Yetman, New South Wales, 12.v.
1977, J. LA. 9/7158) =Comparison with thestype specimen
(IMI 161251) showed that the fungus on this host was con-
specific with D. tripogonis. No other collections have been
made, although 7. lolttformis has been examined at three
other sites in Queensland (Bunya Mountains, Cooroy, Peregian
Beach)® Herbarium specimens Ofpthisespecrcsen lebki mi Lon
many different localities in eastern Australia, were also
searched. No infections Of the, int lorescenceswerescetececd.

Single-conidium isolates from the collection on
T. lolitformts were paired. A teleomorph referable to the
genus Cochltobolus developed in these paired cultures.
Single-ascospore isolates sometimes formed ascomata when:
paired, but never when grown alone. The anamorph formed by
Single-ascospore cultures is indistinguishable from that
produced by the original single-conidium isolates.

Cochltobolus trtpogonts sp. nov. Figures 5 - 8

Ascocarpi atri, basi globosa 300-590 um diam. et collo
cylindrico 880-2000 x 50-150 um. Asci cylindrici, vesti-
Plales bitunicati, (17/0-410 xml sb als oem. en sScOSpOTde
hyalinae, filiformes, versus extrema angustatae, torsivae,
4-9-septatae, 175-275 x 4-6 um.

BivSeyet ea Cochltobolus ravenelitt (1) Ascocarp x50
(2) Asci x200 (3) Ascospores x320 (4) Conidia from
collections 7957(a) and 7963(b), x320. Isolates from
these collections formed the teleomorph of C. raveneltt
when paired.

Figs. 5 - 8 Cochltobolus tripogonis (5) Ascocarp x50
(6) Ascus x320 (7) Ascospore x320 (8) Ascosporic
chlamydospores x320.

343

Sess

344

In) foli1sZeae=maydte in agarovoachil tt 80 (ee DRIP
123757 sho lLotypuss BRIE 227 3542paraty pus.

Ascocarps black, with a globose basal part 300-590 um
diam. bearing a long, ‘straight or slightly curved cylindrical
neck 880-2000 x 50-150 ym (Fig. 5). . Conidiophores: are not
formed on ascocarp body or neck. Asci cylindrical, vestigial
bitunacates slightly tapéredsnear the basco /U—St0e x al 252
18.5 um (Fig. 6). Ascospores filiform, hyaline; helically
coiled in the ascus and completely filling it, commonly
somewhat loosely coiled in the median section and more
tightly in the extremities, gradually tapered to approx-
imately half the maximum width at each end, 4-7-septate,
175-275 x 4-6 pm Fie ew

Single cells of some ascospores become dark brown and
thick walled, while retaining their original shape (Fig. 8).
These darker cells are regarded as ascosporic chlamydospores.
I am unaware of any other reports of this phenomenon in
Cochltobolus species.

In additional testing, inflorescences of 7. lolitforms
on Sach's agar were used as the substrate in an attempt to
produce greater numbers of ascocarps. Ascomata were formed
on the rachis and florets, but no more abundantly than
previously. In some, a basal pedicel 30-150 um high and
30-58 um diam. was present below the swollen fertile section
OteulCEascocanp.

The teleomorph of C. trtpogonts is distinct from that
of most other Cochltobolus species because of the very long
ascocarp neck. Species such as C. cymbopogonis J.A. Hall
& Sivanesan, C. hawattensis Alcorn, and C. raveneltt, which
also have ascomata with long necks, can be distinguished by
other morphological characteristics of teleomorph and
anamorph.

Evidence supporting recognition of Btpolaris as a
genus distinct from Drechslera will be presented elsewhere.
Part of that evidence relates to D. trtpogonts, for which
a new combination is necessary.

Btpolarts trtpogonis (A.S. Patil §& V.G. Rao) comb. nov.
Drechstlera trtpogonis A.S. Patil & V.G. Rao, 1972,
Transeepy.e er my.cOlceoOC moons 4.

345

Single-ascospore cultures representing opposite mating
types of both Cochltobolus species have been deposited in
the culture collection of the Commonwealth Mycological
Pasertutes (I Mim255052mande255655,"0e ravenel77;) JIMIe 235294
Bide 259295 68C. Lr tpodcnts):.

ACKNOWLEDGEMENTS

I- am grateful to Dr R.F.N. Langdon (BRIU) and
Prep eemoutron a IMijeeromercidingmspecimens in their Keeping,
and to Mr R.G. O'Brien for assistance in obtaining some of
the Btpolarts ravenelizt collections.

REFERENCES

Alcorn, J.L. 1978a. Two new Cochltobolus species. Trans.
Demeny CO.) O Cis Ung Ol O07,

Alcorn, J.L. 1978b. Setosphaerta monoceras sp. nov.,
ascigerous state of Hxserohtlum monoceras. Mycotaxon 7:
411-414,

Castellani, E. 1951. Una nuova specie di Cochltobolus.
MY.cOpDa claeMy CO law app lg hOceeo oD fas

Hebert, T.T. 1971. The perfect stage of Pyrtcularia grisea.
Phytopathology 61: 83-87.

Luttrell, E.S. 1976. Ovarian infection of Sporobolus
potrettt by Bipolarts ravenelit. Phytopathology 66: 260-
268.

Luttrell, E.S. 1977. Correlations between conidial and
ascigerous state characters in Pyrenophora, Cochltobolus
and Setosphaerta. Rev. Mycol. 41: 271-279.

Luttrell, E.S. 1978. Biosystematics of Helminthosportum:
impact on agriculture. In Btosystematics in Agriculture.
Beltsville Symposia in Agricultural Research 2,

Bcdlm..AveRombercer, RH. bootes, LV Knutson, GP. l. Lentz,
pp. 193-209. Allanheld, Osmon §& Co., Montclair, New
Mersey.

McKenzie, F. 1968. A study of false smut disease of
Sporobolus spp. M. Sc. Thesis, University of Queensland,
Brisbane.

Patil, A.S. § Rao, V.G. 1972. <A new species of Drechslera
frome nda.) || Fans soi eely COlemoOC. oUt a odo

MYCOTAXON

Vodr eax Le NOs. 8DD ae 40-545 July-September 1981

A PSYCHOTROPIC FUNGUS IN NEPAL

Robert F. Schroeder
Continuing Education
University of Washington
Seattle, Washington

and

Gaston Guzman
Escuela Nacional de Ciencias Biolégicas
Instituto Politécnico Nacional
México, D.F.

SUMMARY

The presence of a Psilocybe species in Nepal which is
close to P. cubensis (Earle) Sing. or P- subcubensis Guzman
and of known psychoactive properties is discussed. This is
the firstyreport or Pagbluingypsilocybe! ingtnesymal lLandadsra.
Speculations on probable early cultural use of this and
other Psilocybe in South) Asiagis,a!1so, presentea.

During 1978 and 1979 Schroeder have been working as
an applied anthropologist for Rockefeller Foundation in
agricultural development in Nepal. On a field trip to
Pokhara in early 1978 he became aware of a bluing species
of fungus. Collections of the species were made in the Pok-
hara valley over a two year period. Although actual speci-
mens were not sent to Guzman, for customs problems, the
Nepal species was positively identified by him as either
Psilocybe cubensis (Earle) Sing., or Psilocybe subcubensis
Guzman on the basis of a set of color photos of the collec-
tions made by Schroeder. These two species are both bluing
and psychoactive; both exist in Mexico, and both are used
by Mexican vIndians. They differ in that P.| cubensis Nas
larger spores, and P. subcubensis has smaller spores, as
discussed Guzman (1978). On the other hand, P. cubensis is
more common in subtropical regions, whereas P. subcubensis
occurs commonly in tropical regions.

This appears to be the first documented report of a
psychoactive Psilocybe species in mainland Asia, except the
reports from Sri Lanka (Ceylon) by Berkeley and Broome in
the last Century, which refer to three species of Psilocybe
(These were recently studied by Guzman in preparation of a

347

monograph on the Psilocybe). Other species of Psilocybe
have been reported by Singer & Smith (1958) from Java and
Dye imazeki & Hongo (1957, 41969.) (from Japan. Recently Guz-
man & Horak (1978) described six probable hallucinogenic
species from New Guinea, New Caledonia and New Zealand, and
Guzman & Watling (1978) reported four bluing species from
Australia and Tasmania. Guzman (in Guzman & Vergeer, 1978)
reported on Agrocybe tibetensis (Mass. ) Guzman, a species
sescribed by \Masseevas !Psilocybe from Tibet in 1906.

The valley in which the Nepal mushrooms were found
lies at an elevation of about 2500 feet and is free of
frost. The area has a monsoon climate with an average
rainfall of about 140 in. per year. Specimens were most
commonly found in abandoned fields or fallow fields where
cattle and water buffalo were grazed, and in maize fields
that had been fertilized with compost. The mushrooms were
often growing on partially decomposed cow dung, and were
found throughout the year. Peak fruiting occurred following
the pre-monsoon rains in May and June, and the mushrooms
were least abundant during the winter months.

There is no known local use for the mushroom, nor are
psychoactive Psilocybe species known to be used elsewhere
in South ASia. Western experimenters who have ingested the
Pokhara species report experiences similar to those occu-
rring with better known species of Psilocybe- Doseage is
relatively high with some individuals ingesting forty or
more mushrooms.

Wasson and Wasson (1957) described the use of Amanita

“muscaria in Siberian religious ceremonies. Later, Wasson

(1972) proposed that Amanita muscaria was used by Indo
Aryan peoples in the soma ritual, which was central to the
development of Hinduism. We agree with Wasson that soma was
earvneus.e Ii lights of the discovery. of an active Psi locybe
species in Nepal, however, we suggest that this species, or
Bnocher yet, to be, discovered Psilocybe species,” is the most
likely candidate for the soma of the Vedas. If this is so,
and the soma of the Vedas is both widely distributed and
totally unknown in modern South Asia, interesting anthropo-
logical questions arise.

AKNOWLEDGEMENTS

The authors are grateful to Dre. Arthur L.- Welden of
Tulane University for reviewing the manuscript. Guzman
express his thanks to CONACyT for support his researches.

348

LITERATURE CITED

GUZMAN, Ge, 19/78. The species of Psilocybe known from Cen-
tral’ and. South” Americas Mycotaxon™/29225-255.

and R- WATLING, 1978. Studies in Australian Aga-
rics and Boletes I: Some species of Psilocybe. Not.
Roy- Bot. Gard. Edinb. 36: 199-210.

and E- HORAK, 1978. New species of Psilocybe
from Papua New Guinea, New Caledonia and New Zealand.
Sydowia 31: 44-54

and P.P. VERGEER, 1973. Index of taxa’in the ge-
nus Psilocybe. Mycotaxon 6: 464-476.

IMAZEKI, Re and’ TI. HONGO, 1957, Coloured Whlustraticn oL
fungisoh Japan. Hoikusha Publ, OsakaemVoOl- al:

and ra, LeIOIA Ni onwe yi Wianirs tks

SINGER, R-, and A-H. SMITH, 1958. Mycological investiga-
tions on Teonanacatl, the Mexican hallucinogenic
mushroom, II. Mycologia 50: 262-303.

WASSON, V-P- and R«-G- WASSON, 1957. Mushrooms, Russia and
History. Pantheon Books, New York.

WASSON, R-G- 1972. Somae Divine Mushroom of Immortality
Harcourt, Brace, Jovanovich, Inc. New York.

MYCOTAXON

eee a ee ST) es ae
Vols A ly eNOtmEZ. app sir 549-5600 July-September 1981

ASCUS STRUCTURE AND FUNCTION IN COCHLIOBOLUS SPECIES

J. L. ALCORN

Department of Primary Industrtes,

Indoorooptlly, Queensland, 4068, Australta

ABSTRACT

Observations by other authors on the
structure of asci and mechanism of ascospore
release in Cochltobolus cymbopogonts are confirmed.
Similar structural and functional attributes of
asci are found in other Cochltobolus species,
including the generic type C. heterostrophus.

INTRODUCTION

Recently El Shafie and Webster (1980) reported their
observations on ascus structure and ascospore liberation in
Cochltobolus cymbopogonts J.A. Hall & Sivanesan. This novel
method of spore release was discovered independently by the
writer (Alcorn 1980), and previously unpublished material is
Presented here.

REVIEW OF LITERATURE

Luttrell (1965) included Coechltobolus in the
Loculoascomycetes, but noted that asci in this genus appeared
to be unitunicate. The ascus in C. nodulosus Luttrell and
C. homomorphus Luttrell & Rogerson had previously been
described as unitunicate by Luttrell (1957) and Luttrell and
Rogerson (1959) respectively. Cochltobolus was again
Piasoitied~as aloculoascomycete by sLuttrel (3 (1975), but nis
illustration of an ascus from the teleomorph of Btpolarts
micropus (Drechs.) Shoem. does not suggest a typical
bitunicate structure. In contrast, Shoemaker (1955) reported
that vasci of C. Sarivus (1to G Kurib.) Drechs. ex Dastur are
bitunicate. This was visible in young asci with immature

DOU

spores, where a canal filled with cytoplasm penetrated the
endotunica at the apex, but was not visible in mature asc1.
Bitunicate asci have also been reported for C. btcolor Paul

& Parbery (Paul & Parbery 1966), C. mtaket Hino & Katumoto
(Hino & Katumoto 1966), C. palmtvora Rao §& Chaudhury (Rao §&
Chaudhury 1964), and C. sttharamit Reddy (Reddy 1976).

Other species of Cochltobolus have been recorded as having
unitunicate asci. In addition to C. homomorphus and

C. nodulosus, they include C. carbonum Nelson (Nelson 1959),
C. cynodontts Nelson (Nelson 1964a), C. gentculatus Nelson
(Nelson 1964c), C. intermedius Nelson (Nelson 1960b),

C. Lunatus Nelson §& Haasis (Nelson §& Haasis 1964), C. sptctfer
Nelson (Nelson 1964b), and C. vtctortae Nelson (Nelson 1960a).

The primary character used in assigning fungi to the
Loculoascomycetes is the bitunicate ascus (Luttrell 1955,
1973). Bitunicate asci have a double-layered wall,
consisting of an ectoascus and an endoascus. The layers
normally are separable, and function in ascospore discharge.
The ectoascus is thin and inextensible, while the endoascus
is) thick gandiextensi ble... @JuStepriOnetomspOLcmLclcascmrnc
ectoascus splits at the apex, andthe endoascus expands
apically, increasing the ascus length by two or three times.
Spores are discharged through an apical pore in the
endoascus. Although many loculoascomycetes follow this
pattern of ascus dehiscence, variations have been recorded.
INNS OMESSPDECLeS wanCircumci Soil eesp illite OCClMomt IEtiIC
ectoascus below the tip, allowing the apical section to be
thrown off as a thimble-shaped cap. This phenomenon has
been observed in Sporormta, Leptosphaerta, Lecantdton, and
Stomtopeltts (Luttrell 1951).

Bitunicate asci lof many species have, an indentation ac
the apex, of the endoascus, formmngea short extension oimeune
lumen filled with cytoplasm (Luttrell 1973). This was noted
in C. sativus by Shoemaker (1955), but is not evident in
other authors' illustrations or descriptions of various
Cochltobolus species. Characters correlated with the
presence of bitunicate asci are ascostromatic ascocarps and
pseudoparaphyses, and Luttrell (1965, 1973) may have relied —
on these when assigning Cochltobolus to the Loculoascomycetes.
Barr (1979) included Cochltobolus in the new family Pyreno-
phoraceae, together with Pyrenophora, Setosphaerta, and
Pseudocochltobolus. She gave no data concerning ascus
structure other than to’ state that asci of fungi in thi's
family are bitunicate.

Drechsler (1925) reported that spore discharge in
Cochltobolus heterostrophus (Drechs.) Drechs. was preceded
by a 'swelling of the ascus and circumcissile rupture in
the apical portion of the ascus wall". In this species
ascospores discharge simultaneously, with sufficient force
for them to clear the ascus in water mounts (Drechsler
1925). Drechsler did not describe or illustrate a cap
comprising the section of the ascus above the split, but
his description of rupturing could indicate that a cap was
formed. Shoemaker (1955) reported that asci of C. sativus
mounted in water commonly dehisced by a circumcissile apical
rupturing, but that they also occasionally broke open at
the centre and base. He obtained no evidence of an endo-
tunica functioning in spore discharge, nor was a residual
endotunica detected in asci which had released spores.
BuuGreling) oo) BLOUNGaLidtaln water mounts, aScl Of: c.
nodulosus split across the apex and the ascospores oozed
out, uncoiling as they emerged. A similar description of
spore release was given by Luttrell and Rogerson (1959) for
C. homomorphus. Discharge of ascospores through a split
in the ascus wall has also been reported for C. carbonum
(Nelson 1959), C. cynodontts (Nelson 1964a), C. gentculatus
(Nelson 1964c), C. tntermedtus (Nelson 1960b), C. Lunatus
(Nelson §& Haasis 1964), C. sptetfer (Nelson 1964b), C.
vietortae (Nelson 1960a), and C. sttharamti (Reddy 1976).
Dastur (1942) recorded that ascospores of C. trvtict Dastur
escaped through an opening formed at the ascus apex by
dissolution of the wall.

Hall and Sivanesan (1972) coined the term 'vestigal
bitunicate' to describe ascus structure in C. cymbopogonis.
With regard to the ascus in this species, they wrote that it
was ‘typically bitunicate but on rupture no extension of
the inner wall vhas been observed’) “Dehiscence was* not
further described but apparently it occurred in the ascocarp,
because one of their illustrations shows ascospores emerging
from the ostiole in loosely coiled groups. Hall and
Sivanesan examined the type collections of C. nodulosus and
C. homomorphus and reported that these species also had
vestigial bitunicate asci. El Shafie and Webster (1980)
found that asci of C. cymbopogonts and C. kusanot (Nisik.)
Drechs. ex Dastur are partially bitunicate, with the endo-
tunica represented by an apical cap which functions in
ascospore liberation. My studies of this phenomenon includ-
ed species not examined by El Shafie and Webster, and this
additional information is presented here.

Joe

OBSERVATIONS

Asci from living material in cultures were examined.
Mountants and stains used in examining ascus wall structure
included water, lactofuchsin, ammoniacal erythrosin, Lugol's
iodine, KOH-phloxine, ammoniacal Congo red, and aqueous
azure A. For studies of ascospore discharge, ascocarps
were crushed lightly in a drop of water to expel asci. In
addition, some ascocarps were suspended individually in
water on excavated slides.

The ascus in Cochltobolus heterostrophus

ThiSesPeC1LeS als, thercenecricatypes(Urechs Letel9 a4) amait
most preparations, the ascus apex did not exhibit a
bitunicate structure. There was commonly a thin unstained
wall 1-2 um thick enclosing the deeply stained cytoplasm.
Even when ascospores did not push completely into the apex,
there was generally no evidence of an apical identation.
Only two asci with a structure resembling an apical cyto-
plasmic channel were seen, one in KOH-phloxine and the other
in azure A. The complex apical apparatus which occurs in
unitunicate asSciswaspnot jseen inethisespecies.

The observations of Drechsler (1925) on ascospore
discharge in C. heterostrophus could not be confirmed. In
water preparations of crushed ascocarps, no discharge of
Spores, from-asciistulieattachedsbasallyetcy cenit tunmerr sso
was detected. » invascis lyinosirec inetne: mountesspore
discharge from the apex was not observed. Some ascospores
escaped, through the base of the ascus, the spores emerging
singly through a narrow aperture at the base of the pedicel.

When unbroken ascocarps were immersed in water, asco-
spore discharge occurred after one hour at approximately
25°C. The released ascospores, sometimess were coiled
together, suggesting that each ascus released its contents
as a unit under these conditions. Empty asci with a split
at the apex were observed still attached to the base of the
locule when these ascomata were crushed. In the same slides
there were short, hyaline thimble-like structures measuring
ca. 30 x 12 um, with a rounded apex and a truncate base.
After pressure was applied to the coverslip, a third, type
of structure was sometimes seen. It consisted of the apical
portion of an ascus, freed by a circumcissile rupture of the
wall. The released apical portion was double-walled. In

bos

some instances the inner wall of this structure became
partly everted when the apical section of the ascus was
forced off. The inner wall was still attached to the outer
wall near the base of the ‘detached structure (Fig. 2). Two
small dark spots were visible at the apex of the inner wall.

The ascus in other Cochltobolus species

Stained preparations revealed no evidence of typical
bitunicate ascus structure in C. cymbopogonits, C. hawattensts
Nicorm, C.exusanot, C. raveneli7 Alcorm, and CG. tripogents
Alcorn, but ascospore discharge in water mounts was seen
HOnscach Species.

My observations on spore discharge in C. cymbopogonts
agree with those of El Shafie and Webster (1980).
Ascospore release begins within 20 seconds when ascocarps
are Crusnedsinewater. slhesspoves are released as aoroup,
ener cine caprdivachrough thevapex of the: ascus, (Fig. 3)”
Each ascospore bundle has a hyaline, cylindrical, thimble-
Pikeestructuresover therapical portion. —lhe) term) ‘endoascal
Caps smuscumbelow ato rereracomtnese: Sstuructures.

The endoascal cap generally separates from each asco-
Spore group as soon as spores are clear of the ascus.
Straightening of the reflexed ascospore tips, and expansion
Opmethe sascospore sheathssand any matrix Lert in thercap,
may be involved in this process. Occasionally the cap
remains ensheathing the ascospore tips after discharge. Asci
which have discharged spores commonly have an irregular
apical split through which the spores were forced. In many,
ClessSp UeawasmmMoOre sO0e1LeSS say eashaped sandecgave, thesascus
apex a) Dilabiate appearance. A broader, less regular rupture
occurred in others, leaving a wide aperture. The torn wall
sometimes was recurved at the margin of the opening.
Occasionallysathevsplit: occurred an) a lateralsposition just
below the rounded apex of the ascus, but this could be an
artefact caused by pressure when applying the coverslip.

The endoascal caps carried out of the asci by the
emerging spores are cylindrical, or slightly widened towards
the base, hyaline, 75-100 x 15-20 um with a smooth rounded
apex and an irregular base which gives the appearance of
having been torn away from a supporting structure. For
comparison, intact mature asci of C. cymbopogonts measure
206-340 x 17.5-20 um. Each cap has a wall up to 5 ym thick

354

at the apex, becoming progressively thinner towards the
base where it is almost indistinguishable. The lumen
contains cytoplasm. “Ati the apexsot the cap gthere sis
sometimes a configuration similar to that seen at the apex
of a typical bitunicateé ascus ,\ that is." aacytoplasm- filed
POLre projecting into thenapicay walle kro 4) eee re para tons
stained in lactofuchsin still had endoascal caps visible
after three months, whereas the mucilaginous ascospore
sheaths dissolved rapidly in this medium. The wall of each
cap had narrowed to approximately half the thickness seen
in’ WaALEY mounts, ,CSpecially=ataciic sapex es hers boeciCccsan
these structures in lactofuchsin strongly suggests that
they are not mucilaginous in composition.

Asci undergo a reduction in length and width after
spore release. Thirty-five asci which had discharged
ascospores measured 185-258 um (mean 220 um). The same
number of apparently mature asci still containing spores
measured 206-330 um long (mean 271 um). Width of empty
asci was 14-17.5 um, and of mature asci 17.5-20 um.
Measurements of ascus length before and after spore discharge
were obtained: individually for five asci. The lengths
were 299 um (before) - 206 um (after), 288-216 um,
263-196 um, 330-242 um, and 319-237 um.

Mature ascomata of C. cymbopogonts liberate asco-
Spores in the absence. of free water. In Petri dish cultures,
long straw-coloured cirri up to 6 mm long and 80-150 um
thick were formed when spores exuded from the ostiole.
These cirri contained ascospores, but no asci, lying
parallel. to the long axis of the cirrus. 7 Ihessporesawere
Straight except at the tip, where many were curved to more
OTA Less uincinate. ~Recently stormedeci ram pod leoreccd
rapidly when mounted in water, but this capacity was lost

Fig. 1. Ascocarp of Cochltobolus eymbopogonts, from culture
of IMI 130402%on Sach's agary+ maize /lLeahe(xG0) serie les.
Broken ascus tip of C. heterostrophus showing partly
everted endoascal wall and darkened tip of endoascus
(x650). Fig. 3. Ascus of C. cymbopogonts with apical
Split through which the endoascal cap and ascospores are
discharged (x650). Fig. 4. Endoascal cap of C. cymbopogonts
(x800). Fig. 5. Ascospore bundle and endoascal cap
of C. trtpogonts (x400).

356

in older, ‘dried cirri, In addition: to “ascospores? the
cirri contained numerous endoascal caps identical to those
produced when asci dehisce in water. Ascocarps with long
CirrilSt1/ 1 had@asciecontainine pascosporessaia tiem lopUse
ascogenous section. Empty asci with split apices were
also present in the ascogenous locule of ascomata from
which ascospores had been liberated. Spore release
probably, occurs ini this part or the ascocarpys themasco-
spores being pushed up the long neck as other asci dehisce.

A discharge mechanism similar to that described above
for C. cymbopogonts was also observed in C. hawattensts,
Cee kueavot, Ca ravenel1c,. and (6 mitnipogonis (asl nesendcascal
caps in these species are similar in appearance to those
of C. eymbopogonts (Fig. 5).

An hypothesis to explain ascus structure and function in

Cochltobo lus species

It is postulated that the ascus in Cochltobolus is
bitunicate, and that the endoascus is thickened only in
therupper portion. iit issturtherssucgestedmthacscie
mechanical strength of the endoascus in the lower part is
not @reat, vand that | thisevesults gin ithemwalleotete
endoascus rupturing in a circumcissile manner under the
force exerted Dy sescapinomascospores,.

The presumed events up to and including spore discharge
are»as follows.» “As ascospores mature, theyascus@us
stretched to accommodate them. Eventually the ectoascus
Splits cat thetap..) Ihe endoascusemay expandyasminga mormal
bitunieate ascus (at this*stage. Its consideredemore
likely, however, that the endoascus is forced out of the
ectoascus by the expanding ascospore bundle and ruptures
at a point approximately one third of the ascus length from
themgapex.7 sIhevendoascal capeis=thenscarrlcdaawavedased
sheath’ over the tips, of sthesascospores.. Angalcemativesto
this hypothesis would be that the lower portion of the
endoascus is firmly bonded to the ectoascus, and cannot
expand to accommodate the increase in pressure which
precedes ascospore release. “Jf present, thesendoascus in
this lower region is too thin to be distinguished by light
microscopy. It might also be argued that the endoascus
is not complete, being present only in the upper third of
the ascus. This seems a less plausible explanation, because
it would require the endoascus to be attached to the
ectoascus in a narrow region in the upper part. If both

So,

walls were present in young, shorter asci, basauxic growth
of the ectoascus would have to occur to bring about the
condition of mature asci. Transmission electron microscopy
possibly would help resolve these uncertainties concerning
ascus Structure.

Note on ascocarp morphology in C. eymbopogonts

The reports of Hall and Sivanesan (1972) and El Shafie
and Webster (1980) indicate that ascomata of C. cymbopogontis
are sessile, with a rounded ascogenous basal portion and a
long cylindrical neck. _ When grown on Sach's agar + leaf of
Zecemaycklny the type culture ot this especresmGlMis130402)
produces ascomata in abundance. Erect, cylindrical black
sunOMAtambirStadevelop onetnicsleatapleces, sespecially
along the major veins. The ascomata differentiate singly
auethewapex of each Stroma, =sOuthaveat Macurity each
ascoma has ascylindrical’stérile: base (rigs 1). These
bases measure 500-1400(-1700) x 175-260 um. On water agar
+ wheat straw, ascocarps are less numerous and most are
sessile. Presence of a sterile stromatic base was one of
the characters used by Tsuda, Ueyama, and Nishihara (1977)
to distinguish the new genus Pseudocochltobolus. Clearly
this facet of teleomorph morphology in C. cymbopogonts
is conditioned by substrate. If this also applied in other
species, it would be unsuitable for use as a generic
eraee rion

DISCUSSION

Roguseno tac Loarawieticmechemuypemspecies) Of
Cochltobolus was studied by either Luttrell (1965, 1973) or
Barr (1979), but both authors assigned this genus to the
Loculoascomycetes. Barr stated that asci are bitunicate
in the family Pyrenophoraceae, in which she included
Cochltobolus. Luttrell (1965) believed that asci are
unitunicate in Cochltobolus, while Hall and Sivanesan (1972)
proposed the term 'vestigial bitunicate' to describe ascal
structure in this genus. The data presented here and by
El Shafie and Webster (1980) support the contention that
Cochltobolus should be classified in the Loculoascomycetes,
and offer some explanation for the differences of opinion
concerning ascus structure in the genus. It is suggested
that the term 'vestigial bitunicate' used by Hall and
Sivanesan should be retained, because it seems that the
endoascus plays no active role in spore discharge from the

358

ascus. In this respect my interpretation of ascospore
release in Cochltobolus differs from that of El Shafie and
Webster (1980). They referred to a protrusion of the
endoascus enclosing the spores, followed by spore release.
I believe the endoascus plays a much more passive role,

and that it is forced out of the ectoascus by the expanding
ascospore mass. However the process occurs rapidly and
individual steps cannot be distinguished.

Unitunicate asci may have an outer, denser layer which
is not structurally different from the inner wall layer
(Grifriths 1973S) One olathe interpreta tionsaoieascus
structure in C. cymbopogonts and similar species raises
the possibility of fusion of walls in the lower part of the
ascus. If transmission electron microscopy shows this to
be correct, then the ascus in Cochliobolus may represent
an intermediate form between unitunicate and bitunicate
GY pes:

Five of the Cochltobolus species in which ascospore
discharge was observed have narrow cylindrical asci. Fresh
material of other species such as C. sativus and C. vtctortae,
in which asci are much broader, should be examined to
ascertain whether the discharge mechanism is the same. It
is also important that ascus structure in Cochltobolus be
observed by transmission electron microscopy, to relate
ultrastructural detail to form and function as interpreted
after examination by light microscopy.

ACKNOWLEDGEMENT

Pethanke Dre wane See Ontons mt Oresthemty pescuLtLuresot
C. eymbopogonts, and Mr. A.E.. El Shafie'for a culture of
C. kKusanot.

REFERENCES

Alcorn, J.L. 1980. Generic concepts in Drechslera sens. lat.
and correlated teleomorphs. Ph.D. Thesis, University
of Queensland, Brisbane.

Barr, M.E. 1979. A classification of Loculoascomycetes.
Mycologia 71: 935-957.

Dastur, J.F. 1942. Notes on some fungi isolated from 'black
point' affected wheat kernels in the central
PYOVINCES. Geel ndhe) Gash Cwm Gln ee noreny, bea a.

590

Drechsler, C. 1925. Leafspot of maize caused by Ophtobolus
heterostrophus,n. sp., the ascigerous stage of a
Helmtnthosportum exhibiting bipolar germination.
rea Or Ce cReCShayo lice (01-776.

Drechsler, C. 1934. Phytopathological and taxonomic aspects
of Ophiobolus, Pyrenophora, Helmtnthosporium, and
a new genus, Cochltobolus. Phytopathology 24: 953-
SIS,

Ele ondiie mA cb anGaN CDS tcl a) ml GUNmmmASCOSDOLCm iberatilon
in Cochltobolus eymbopogonis. Trans. Br. mycol.
SOCEay/ Ocml4 ie 146),

Griffiths, HB. 1973. Fine structure of seven unitunicate
DY renomy ceterascis. trans.) Din emycOl.Wo0C.. 00.201 —
Zila

Halipmue Nee Geol vanesan,eA, 19) 28 COCiLTODOlLUuSsastaLe OL
Curvularta eymbopogonts. Trans. Br. mycol. Soc.
SO eels

Hino, I. & Katumoto, K. 1966. Notes on bambusicolous
DUC lee (cy ete ieee) tee DO Umea co oe ie

PuGtre Leet ores! OS mel axOnonye Oe Leh yrenomyce ves:
Univ. Missouri Studies 24(3): 1-120.

mut relies 1955.5 Ihesascostromatic, Ascomycetes:
Mycologia 47: 511-532.

Luttrell, E.S. 1957. Helminthosportum nodulosum and related
species. Phytopathology 47: 540-548.

Pitre ol 905 Classi-f)catlOneOretne Loculoascomycetes.
Phytopathology 55: 828-833.

Luttrell, E.S. 1973. Loculoascomycetes. In The Fungt, an
Advanced Treattse, Vol. IVA, ed. G.C. Ainsworth,
F.K. Sparrow & A.S. Sussman, pp. 135-219. Academic
Press, New York.

Luttrell, E.S. § Rogerson, C.T. 1959. Homothallism in an
undescribed species of Cochltobolus and in
Cochltobolus kusanot. Mycologia 51: 195-202.

Nelson, R.R. 1959. Coehltobolus carbonum, the perfect stage
of Helmtnthosportum carbonum. Phytopathology 49:
807-810.

Nelson, R.R. 1960a. Cochltobolus victortae, the perfect
stage of Helmtnthosporium victortae. Phytopathology
5037 As4-775:

360

Nelson, R.R. 1960b. Cochltobolus intermedius, the perfect
stage of Curvularia tntermedta. Mycologia 52:
7752775.

Nelson, R.R. 1964a. The perfect stage of Helminthosportum
eynodontts. Mycologia 56: 64-69.

Nelson, R.R. 1964b. The perfect stage of Helminthosportum
sptetferum. Mycologia 56: 196-201.

Nelson, R.R, 1964c. The perfect stage of Curvularia
gentculata. Mycologia 56: 777-779.

Nelson, R.R. & Haasis, F.A. 1964. The perfect stage of
Curvularta lunata. Mycologia 56: 316-317.

Paulo TAR. GeearDery, slo GeeloOG mm ne DeChLecums tatemor
Helminthosporium btcolor. Trans. Br. mycol. Soc.
49: 385-386.

Rao, P.N. & Chaudhury, R. 1964. A new species of
Cochltobolus from Hyderabad - India. Mycopath.
Mycol. apple 25:7 36-387

Reddy, S.M. 1976. A new species of Cochliobolus. Indian
Phytopath. 29: 199-201.

Shoemaker, R.A. 1955. Biology, cytology, and taxonomy of
Cochltobolus sativus. Can. J. Bot. 33: 562-576.

Tsuda, M., Ueyama, A. & Nishihara, N. 1977. Pseudocochiobolus
ntstkadot, the perfect state of Helminthosporium
cotets. Mycologia 69: 1109-1120.

MY COTA XON

DOs aN Ll NORS Ee. sD pearoO 1-566 July-September 1981

A PRELIMINARY DISCOMYCETE FLORA OF MACARONESIA:
PART 6, GEOGLOSSACEAE*

RICHARDS Pas WORE

Plant Pathology Herbarium, Cornell University
Ithaca, New York 14853 USA

"Was never eare, did heare that tong."

Edmund Spenser
AN ELEGIE, OR FRIENDS PASSION, FOR HIS ASTROPHILL, Line 110

Order HELOTIALES
Suborder HYMENOSCYPHINEAE
Family GEOGLOSSACEAE Corda 1838

KEY TO THE KNOWN MACARONESIAN GENERA

a INSCOSPOLCSmaD GOW amd eel a (UL LUY gee tere lores coerced se oct Ges 2
Me SCOSDOLCCMIIVALING esa eMALULILY. so atetee je site oe eee es ertie 4,
2(1). Hymenial setae present. Trichoglossum
2a rem ymeniae Setde: —aDSeN tures. teeta’. <cpctete) <fs 3
Bi2') 3 PatapiycesmeeseniGtedestOnsiyMmen UM note GUM G
down stipe; stipe dry. Geoglossum
3'(2'). Paraphyses running down the stipe; stipe glu-
tinous. Gloeoglossum

4(1'). Ascoclavula black or blackish.
Thuemenidium

Lytle) eA SCOClAVUlas paler, note Diackishs
Microglossum

GEOGLOSSUM Gillet 1879

One known Macaronesian species

* The parts of this flora will appear in irregular order.
Reprints of individual parts will not be available for dis-
tribution.

362

ihe Geoglossum umbratile Saccardo, Michelia 1: 444.

1878, var. umbratile

MISAPPLIED NAME: Geoglossum
nigritum (Rens samba mNG Ken

RECENT TAXONOMIC TREATMENTS:
Benkert (1976), Maas Geesteran-

us (1965), Mains. (1954), JNann—
teldtmigou2)

PREVIOUS MACARONESIAN RECORDS:
None.

TYPE LOCALITY: Italy.

KNOWN MACARONESIAN DISTRIBUTION
CANARY ISLANDS.
Tenerife. CUP-MM 21 (OSC, TFC,
to™ be -issueds-in.] Kort 4&
Gruulif aL 1SC en Ocoee aie ie
CLEG):. 12798e ChE C Ame eOS
(el ase

SUBST RAGA == (On seSOl leet pemecdil Che Samia imcdenon
roadside banks.

Notes: Mains (1954) and Nannfeldt (1942)
use the name G. nigritum, and our Ma-
Garonesianm collection S Meweres imiilallyemi.
dentified as G. nigritum var. nigritum,
but Maas Geesteranus (1965) has shown
that name applies correctly to a basidio-
mycete (Persoon originally described it
dSeyClaVariatenloni ta, aed imate acmabccn
consistently misinterpreted since then).

GLOEOGLOSSUM Durand 1908

G. umbratile
v. umbratile,
3 ascospores,
CUP-MM 1272,

aLOCOR

One known Macaronesian species

1. Gloeoglossum glutinosum (Pers. : Fr.)

Mycol. 6: 419. 1908.

1827.

= Geoglossum lutinosum Pers.,
LI ah 90 eee eS a oy St.

Durand, Ann.

Obs... .Myc. us
Myc. 1: 489.

RECENT TAXONOMIC TREATMENTS:
Mains (1954), Nannfeldt (1942).

PREVIOUS MACARONESIAN RECORDS:
None.

ley PL OCAM TTL Y<@Europe.

KNOWN MACARONESIAN DISTRIBUTION
MADEIRA.
Madeira. CUP-MM 1566.
CANARY ISLANDS.
Gomera. CUP-MM 1168(0O).
Tenerife. CUP-MM 1187(TFC).

SUBSTRATA: On soil.

Notes: Though most modern authors
Come nOtmsacCeDtmDULA TI CmSmmseCrCOalc
genus, the glutinous ‘stipe makes it Gl

an eas taxon to recognize even in glutinosum, 1
on > a immature asco-
the field.

Spore, sou mature,
CUP-MM 1566,
xn OOO:
MICROGLOSSUM Gillet 1879

One known Maraconesian species

1. Microglossum olivaceum (Pers. : Brioeeo letme lam pr
inmenvean Wicca) MNes ali Yoo Nevis)

RECENT TAXONOMIC TREATMENTS: Dennis (1978), Mains
(1956), Nannfeldt (1942).

PREVIOUS MACARONESIAN RECORDS: None.
mY Ere LOCA hy see urope.
KNOWN MACARONESIAN DISTRIBUTION
CANARY ISLANDS.
Tenerife. CUP-MM 1307(TFC).
SUBSTRATA: On soil.

Notes: Mains (1956) included Thuemenidium (= Corynetes)
in his very broad concept of the genus.

ID
Tea

ay a

M. olivaceum, 6 asco-
SPOCGS, a Amea SClee WA +
Pole wiwallechanne 16
ascospores, CUP-MM 1307,
xe L000:

Th. atropurpureum, mature ascus
with J+ pore wall channel, immature
ascus, 12 \ascospores, CUP=MM 1371;

x LOO:

THUEMENIDIUM Kuntze 1891 emend. Maas Geesteranus 1964
One known Macaronesian species

1. Thuemenidium atropurpureum (Batsch : Fr.) Kuntze,
ReEVisS. 3 OC Dil wees) el oo e.

= Corynetes atropurpureus (Batsch : Fm) Dur=
and. ‘Anne Mycol. 86:4] 42161 908"
= Microglossum atropurpureum (Batsch : Fr.)

Karst Acta. Soc shatindgw hl. Leen. .o1n
LiOe 1885.

365

RECENT TAXONOMIC TREATMENTS: Dennis (1978), Maas
Geesteranus (1964), Mains (1956: Microglossum), Nann-
feldt (1942: Corynetes).

PREVIOUS MACARONESIAN RECORDS: *Bafiares Baudet &
eee 1980), nt Beltran, Tejera (1980),**Cool (1924), ***Cool
(1925).

Pees OCA TRY eEurope.

KNOWN MACARONESIAN DISTRIBUTION
a CANARY “I1SUANDS:
*Gomera. *CUP-MM 1371(TFC|[=TFC 770])).
xx ***Tenerife. CUP-MM 1136(0), ** ***1652(L).

SUBST RATA: On soil:

Notes aaa NOuGns sthesechilerion. Of Malleamorphous,. colored
pseudoepithecium has been used to distinguish Thuemeni-
dium, it is often absent. Mains (1956) I think ‘correctly
Synonymized 3 species Durand (1908) distinguished in
has’ monograph, Corynetes atropurpureus, C. purpuras-—
cens, and C. robustus, the last with an epithecium

"lacking or inconspicuous," and with more robust asco-
mayulayswand= “somewhat, donger » spores. The) collection
Bronte, Gomera, — ceporied by iBanaress Baudet <Sival-seswas

Baken during onesof our expeditions to; Gomera, together
Sptumeticerstatisor them Universidades de ‘LagiLaguna... It
Paice NOn trace oim a scolored = pseudoepithecium and? thus
mronches sO. robustus, and valso, hasevery large ascoclavu—
la, but the ascospores are within the typical range of
PeeearvopunpuULeum. | A= better criterion = sperhaps, is) the
peculiar dual-hyphal structure in the stipe, evident in
the Gomera collection as well, which was stressed by
Maas Geesteranus (1964). Mains (1956) included this in
Microglossum, while Maas Geesteranus points out that
it may belong in Geoglossum, differing mainly only in
the colorless (and non-septate) ascospores. Corynetes
arenarius =(Rostrup) Durand; from Greenland, is’) talso
a Thuemenidium, but C. globosus (Sommerf. : Fr.) Dur-
and from Scandinavia, the Alps, Greenland, and proba-—
bly Canada is a Sarcoleotia according to my studies,
though treated in Corynetes by Nannfeldt (1942); Maas
Geesteranus (1964) showed it could not be a Thuemenidi-
um.

—_—

366

TRICHOGLOSSUM Boudier 1885

One known Macaronesian species

1. Trichoglossum hirsutum (Pers. : Fr.) Boud., Hist.
classific., discomyc., Europe, sip 8'c0.18190/,0) Van.
hirsutum

RECENT TAXONOMIC TREATMENTS:
Dennis (1978), Mains (1954),
Nannfeldt (1932).

PREVIOUS MACARONESTAN RECORDS:
*Baagge & al. (1972), ** Beltran
Se Wildpret (19/5)G0 1 Beltran lejera
(1980).

TY Pre pOGAL TRY rE Urope.

KNOWN MACARONESIAN
DISTRIBUTION
*MADEIRA.
*Madeira. CUP-MM 1492(TFC,
CeCe WAS, oi tisytem ioyas
PACER Me Virpe ZAS GIA VAG)
(Cea Sai Spi) eee) Ce
**CANARY ISLANDS.
La Palma. CUP-MM 668, 669
GhEG= OSE).
**Tenerife. CUP-MM 1113(0),
LWA) LUZON 1a 6i/
(Oe L26UCT EG 4 1296,
1Z29SECTECG, LOSCe to “besus—
sued = int) @DiSc.s eh xs sn)
**2507(TFC), **2509(TFC).

ae EF ie Ee Oe Se

SUBSTRATA: On - bare soil and
among mosses.

Tr. hirsutum var. hirsutum,
2 15-septate ascospores,
ascus japex jin iodine,
CUP-MM 1113; 7-septate
ascospore, CUP-MM

2007 all, x 71000;

MYCOTAXON

Ore LiitleawNO arcem DDO 07" 250 6 July-September 1981

The Psilopezioid fungi. VII.
Aenewespeciess Ofars1 Lopez tas from sPrance.

DonaldpHs) pirster.
Farlow Reference Library and Herbarium of Cryptogamic Botany
Harvard University, Cambridge, MA

and

Frangoise Candoussau
22, rue Ho6-Paris, 64000 Pau, France

Psilopezia nummularialis Pfister and F. Candoussau sp.
nov.* Receptacles gregarious, 1-3 cm diam, sessile, convex,
later convoluted, broadly attached to the substrate, sur-
rounded by a white mycelium which is particularly evident
mimanvedsmateria ls) Externalesurtace lighter.) sHymenium
gazelle colored, a rich ochraceous, the pigment soluble.
The flesh paler than the hymenium 1 to 2 mm thick toward
the center, translucent, brownish when dried.

OULCracxGipu bumeotetwomind tstanc taelayonms wmainnerslayer
Peeteparallel septate thyphacawhiche becomesloosely yinter-
fwined, 9-10 jumein diam onethe sinner position sand, 20-35 °pm
POWwALGe CHC OULCTS Ones ObEtwoRcel |S. 01 stnissinner layer.) The
outer layer, non-gelatinous, of interwoven hyphae about
12-15 pm in diam with a few free hyphal tips which sparsely
Bover the outerysuntace .

Medullary excipulum of textura intricata of cells
Poo um in sdiam.

ASeCimc-spored ,=400 0x 18-22eumsbroadgwith evident
SrOZ1ers..

Paraphyses encrusted with brown amorphous material,
Pranching and septate below 9-11 pm.

HOLORY BE aes metroucede scald xepOUnT awa cerre mendrort
inonde periodioquement, avec Pachyella sp. et Climacodon
pulcherrimus (Berk. §& Curt.) Nikol. Forét de Bugangue
(Oloron) Pyrénées Atlantiques. F. Candoussau. 14.1X.1980.
Holotype see elsotypes F.C. 10. fo7 oe andgeUPes 0876.

Other collections made in October and November in the
same locality are in FH and the collection of the junior
author.

This species is close to Psilopezia nummularia Berk.

*Psilopezia nummularialis Pfister & F. Candoussau sp. nov.

Stmilis Psilopezia nummularia Berk. sed ascosports 26-29 x 12-15 pwn,
ascets 400 x 18-22 ym et hymenits ochracets.
Holotypus: sur trone de Salix pourri, a terre, endrott inonde peridto-
quement, avec Pachyella sp. et Climacodon pulcherrimum (Berk. & Curt.)
Wikol. Forét de Bugangue (Oloron) Pyrénées Atlanttques. F. Candous-
sau. 17.IX.1980. Holotype FH; Isotypes F.C. no. 37571 and CUP 59826.

368

Figure 1. Apothecia of Pstlopezta nummulartalis scale = 2 cm.

which has*been;collected in Europe and ‘1s "known an some or
the literature. under the name \Peziza‘atroviolacea Bres=,
shown to be a later synonym (Pfister 1973). The two
species can be distinguished as follows:

P, nummularialis P. nummularia

See ochraceous olivaceous, or nearly black

color
Asci 400 x 18-22 pm 275-300 x 20-27 um
Ascospores 26-29 x 12-15 pm 29-40 x 14-20 pum
Outer zone

of outer 10-17 pam up to 35 pm

excipulum

In anatomical features Psilopezia nummularialis and P.
nummularia are quite) similar.) slt should be pointed out,”
that a report by Beller (1972) of Psilopezia nummularia is
inetacterachvellarbabang toni ieab erik wiGeb i

The work of the senior author was supported in part by National
Science Foundation Grant DEB 8023018. We thank E. Jarias for photo-
graphs.

Literature Cited

Beller, J. 1972. Psilopezia nummularia Berkeley. Documents mycolo-
gigues., 5% ).1=4¢

Pfister, D.4H.9 719737 sThel psilcopezioid wfungivell Laine genus hcalopes
zia (Pezizales). 60: 355-365,

MYCOTAXON

Oued NO. 25 DD o092572 July-September 1981

NOTES ON SOIL FUNGI ISOLATED FROM A 15-YEAR-OLD ASPEN STAND
IN INTERIOR ALASKA

LOLA K. OLIVER and KEITH VAN CLEVE

Forest Sotls Laboratory, Universtty of Alaska
Fatrbanks, Alaska 99701 USA

SITE DESCRIPTION

The sampling site, a quaking aspen (Populus tremulotdes Michx.)
Stand located 42 kilometers northeast of Fairbanks, Alaska, is an area
between 150 and 210 meters above sea level and is situated at 146°50'W
longitude and 64°54'N latitude. Soil is alluvial silt-loam of the
Goldstream-Saulich series. The lowland site has a slope of between 3
and 7%. Mean annual precipitation and temperature for the Fairbanks
vicinity are 28.7 cm and -3.4°C, respectively. A severe forest fire 15
years prior to sample collection destroyed virtually all surface vege-
tation on the study area. Nearly all trees and other plants now pre-
sent in the area are less than 15 years of age. Present cover is
Suspected to have originated from surviving root stock and sprouts.
Average tree height in the burn area is approximately five meters;
diameters are eight centimeters or less, and ground vegetation is very
sparse.

METHODS AND MATERIALS

Prior to sampling for soil fungi, factorial applications of N, P
and K fertilizers were made to the sample plots as part of an on-going
experiment dealing with organic matter production. The present study
was limited to sampling from the nitrogen, phosphorus, potassium appli-
cation and control plots of that experiment. Plots were fertilized in
the fall prior to sampling and again in May of the same year at the
rates of 112 kg/ha NH4NO3 for the nitrogen plots, 56 kg/ha P90, for the
phosphorus plots and 112 kg/ha KCl for the potassium plots. Sampling
began in late May and continued once a week for eight weeks. A 6.5 cm
corer was used to collect three cores of surface material from each of
two subplots in each of the three replicate fertilizer and control
plots (eighteen cores for each treatment). Cores for each treatment
were combined for fungal isolation. The litter layer of the site was
present in too small a mass to be used alone for sample analysis. The
fermentation and humus layers were indistinguishable; hence the entire
organic layer of the cores was used. The depth of this total organic
layer was never more than 3 cm but never less than 2 cm. Organic con-
tent of the layer was calculated by sample ignition in a muffle furnace
ace/00-C toranourn hours. «it ranged from 4124910, 81/2. »So1l* pH canged
from 4.7 to 7.5 and was determined on a saturation paste using a glass

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4uebe 39euyXa [LOS uadsy = EIS [OuzU0) = 9

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[ebuaq asOu yyLM 4ebe asouzxap 03230 = guvdd (50%) snuoyudsoud = d
Aebe asouzXap 07e30qg = Wad (fonPHN) uabouziN = N

“ds um21thy,01Uu0)y,

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ULWALLLNA SstumBouazay snyouhyiobh7z

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electrode pH meter. Above ground temperature for the study period
ranged from -4°C to 28°C. Soil moisture content over the period of the
study ranged from 12.4 to 176.6% (wet weight %).

Samples were plated in replicates onto four different agar media.
A different agar was used each week. The agars used were: potato
dextrose agar at pH 4.0 (PDA); potato dextrose agar with rose bengal
added (PDARB); potato dextrose agar with rose bengal and streptomycin
added (PDARBS); and aspen soil extract agar (SE). The dilutions from
which the fungi were taken were 10-5 for all potato dextrose agars and
10-4 for the soil extract agar. Five colonies from each of five plates
prepared for each area were isolated and cultured on potato dextrose
agar pending fungal identification.

RESULTS

Species were identified using methods outlined by Raper and Thom
(1946), Barnett (1960) and Gilman (1971). In all, 800 fungi were
isolated. Duplicate species were discarded and by the end of the study
43 species had been isolated. Twenty of the more difficult species to
identify were sent to Dr. J.A. Von Arx at the Centraalbureau voor
Schimmelcultures in the Netherlands for identification. Those that
his staff confirmed or identified are indicated by an asterisk in the
accompanying table of fungi. Fungi not identified were given a number
notation. Four of the Mycelia Sterilia species had noticeable clamp
connections, indicating the presence of basidiomycetes.

ACKNOWLEDGEMENTS
We would like to thank Terrence A. Moore and Dr. Gary A. Laursen
for their advice and criticism of this paper. The research was funded
in part by the McIntire Stennis Cooperative Forest Research Program.
LITERATURE CITED

BARNETT, H.L. 1960. Illustrated Genera of Imperfect Fungi. 2nd ed.
Burgess Publishing Co., Minneapolis. 225 p.

GILMAN, J.C. 1971. A Manual of Soil Fungi. 2nd ed. The Iowa State
University Press, Ames. 450 p.

RAPER, K.B. and C. THOM. 1946. A Manual of the Penicillia. Hafner
Publishing Co., New York. 875 p.

MYCOTAXON

VOL Nite NOs la DOs 75-404 July-September 1981

COPRINUS SECT. HERBICOLAE FROM CANADA, NOTES
ON EXTRALIMITAL TAXA, AND THE TAXONOMIC POSITION OF A
LOW TEMPERATURE BASIDIOMYCETE FORAGE CROP
PATHOGEN FROM WESTERN CANADA

S.A. REDHEAD

Btosystemattes Research Institute, Research Branch,
Agrteulture Canada, Wm. Saunders Bldg., C.E.F.,
Ottawa, Ontarto, Canada Ki1A 0C6é

&
J.A. TRAQUAIR

Agriculture Canada Research Statton,
Lethbridge, Alberta, Canada, T1J 4B1

ABSTRACT

The pathogenic low temperature basidiomycete (LTB)
affecting turf grass, forage legumes and winter cereals
in western Canada is described as C. psychromorbidus sp.
nov. A second western species is described as
C. maysotdtsporus sp. nov. Coprinus section Herbtcolae
is revised and a key to the accepted species given. The
study of type specimens led to the conclusions that
Coprinus satchtae Reid and C. rhombisporus Orton are
synonyms of C. frtestzi Quél. Coprinus brasstcae Peck,
C. melo Favre and C. suburticicola Pil. & Svr. are
synonyms of C. urtictcola (Berk. & Br.) Buller.

Coprinus pseudofriesti Pil. & Svr. and C. xantholepts
Orton are synonyms of C. phaeosporus Karst., and Coprinus
amphibius Anast. is a synonym of C. kubickae Pil. & Svr.

In addition, Coprinus paleotroptcus sp. nov.,
C. neotropicus sp. nov. and C. austrofriesitt sp. nov. are
described for tropical collections previously reported as
C. urttetcola, C. brassteae and C. frtestit.

A previously unidentifiable low-temperature basidiomycete (LTB)
known to cause snow mold damage to turf grasses, winter cereals,
forage legumes and native perennials (Broadfoot 1936, Broadfoot &
Cormak 1941, Lebeau 1969) has been identified as a species of

374

Coprinus in the section Herbicolae (Traquair 1980). The specific
identity of this Coprinus could not be ascertained from existing
publications. Furthermore, although a re-evaluation of type materials
for names published for species here considered in section Herbicolae
revealed some inconsistencies, none of these species were conspecific
with the pathogenic Coprinus. The fungus and a second species
isolated with it are described as new. Descriptions of other species
known to occur in Canada, a discussion on related taxa and a key to
species accepted in section Herbicolae are also given. A description
of the cultural characteristics of one species follows the taxonomic
portion. Three tropical species are described as new and problematic
species are discussed.

A key to Coprinus section Herbtcolae

A. Spores blackish to blackish brown and rapidly losing

their colour and form in concentrated sulfuric

acid solution vor doingtso within J—Seminutese ys: ces se ciee melee s B
A. Spores pale to dark brown or blackish brown,

remaining relatively dark and intact in

concentrated sulfuric acid solution for

over 5 ‘minutes sess ce ols c's oles scovsteiecdse saa cle: eieicre «vile a creretclensiorersl cretetets D
B. Veil elements with hyaline thin walls; spores
blackish? brown) OVOLO weet cestete eters rete cietetcecte stercrete C. kubtekae

B. Veil elements with hyaline moderately thin walls
lightly incrusted with brown pigments; spores
blackish, shaped like maize kernels ........cccccccccseccees (e

Co Sporese5 25-6. 58 X eos Uo ace Nee O—e coms
Imbbgeesa18h A AAD OSA OOO E MOG OI5 ObC0d BOUCHE SOO OOU CECE C. argenteus
GC. Spores 7=9 X46—/7.6' X°5.43—5.3um;
NOL ENPAME DI Calne cic cies es ere lepeiets ciaisters citrate cca choisrers che C. maysotdtsporus
D. Spores globose to subglobose,
4,.5-5.0 X 4.5-4.8um, apically rugulose ........... C. herinktt
D. (opores notagloboseys Larger pa Smooch wat. «sss /cicislciats eieleietetelclats ete E

E. Veil elements hyaline or subhyaline, in one
species tending to become faintly yellowish
with age or on drying stelleloleteletalcletelciclotelcdeleloleteletcletelclencieleretetonetonetolenalelets F
E. Veil elements prominently pigmented, brownish
EO TGO dens DOWN cts ecenie riereiciele oreeretelsteceiere sis oie aiele etetete eterete sreteremerc at ite I
F. Veil elements thick-walled, over
O Sumpthickein places = acc eile clacis canine 4 sites cieicie aca citer aerate G

D715

F. Veil elements thin-walled, under

Ossi na ikelle gad Soyo bed ae Bonn Hoe Meer emnene hea H
Spores 9.6-11.8 X 7.2-8 X 7-9um;
SMALE) CAN Sterectetele ctrise stoiiereteietersisie es clorsme cre lsians atolete ¢ C. austrofrtestt
Spores 7.5-9.3 X 5.5-7.9um;
N. Hemisphere ...... oie iesecu tenet cnerehtichore siattict ate: tonsils etolt ets iste taieks On GRRE
Spores 3.8-4.2 X 4.5-5.3um;
IN SUpAITIS Td CAN ant etetats cootele rsiutelerere:sieitic's s/eiei ciereiers crsie elcichelate scoisie/s Cm DURKLL

H. Spores predominantly ovate in face

view, slightly compressed in profile,

6.1-8 X 4.5-5 X 5.8-6.7um, walls

ChickeninG pBALTACAN wists stccietes crciete iets siete ts crs C. paleotroptcus
H. Spores predominantly broadly ellipsoid

to near subglobose, 5.5-8 X 3.9-5.1lum,

walls relatively thin; N. Hemisphere ........... C. urttetecola
H. Spores ellipsoid to ovoid,

8-9.5 X 6.5-7.5um, walls relatively

CHUTES seAMELIUCANG he's sietatyertetetc sie sea crete hate ete's’s ¢ C. neotroptcus
Veil element walls refractive, thickened,
often 0.5-1.0um thick over long segments;
elements much branched, with some units like
MacuLed edee~eant. Crs; sGOLACH Wine COLOULMMs se ciclete’s stelcicis sie srctaleielcre ears J
Veil element walls thin to only slightly
thickened; elements moderately branched, at
most some units shaped like young blunt deer
cupyelitvesyy leiqe,euucla wig! @eMVelie 5. ooGasqUTUdC GGUUU ROK DOG UOSHOOC On mC L
J. Spores obscurely to prominently angular

in face view, being an elongated hexagon ....... C. herbtvorus
We MEOOOLeS LOVaLeat One LIptical sin sLace LVILGW Wsiclsisiets css clclale¢ siansiatc K
Spores 5-7(-8) X 5-6um, in face view, broadly
ovate, less so in profile; with a nearly lateral
eccentri CRapiculus on Many ;aEULOpeAan wer. secs ele ce ore C. phaeosporus
Spores (6.4-)7.2-8.8(-9.6) X (4.2-)5-6(-7.3) um,
in face view, typically ovate to broadly elliptical,
varying in different isolates, some with nearly all
spores broadly ovate, in others with nearly all
elliptical; apiculus slightly eccentric to nearly
Pacerall seNObtneAnerd Caliitsc\ste's sivcs ccs e siniclaibis = sles C. psychromorbidus

CWA®

K. Spores 9.5-11.5 X 6.5-7.0um, in face view
ovate to elliptical; apiculus slightly

eccentrics Jamaican” (Oniwo0d) io... caiusecas ests eee C. cetnehonensts
L. Spores 5.7-7 X 4.5-6um;

SOUL TAMEPICANE stds oleit cole cleieic.e o opnctmte welere sieteters C. subttgrinellus
L. Spores 8-10 X 6.8-7.lum; European ......e-ceees C. ttgrinellus

SPECIES FOUND IN CANADA

*Coprinus friesti Quélet (1872: 159, tab. 23, no. 5) Figs. 13,14,16,
29 ,34,44-47.
= Coprinopsts friestt (Quél.) Karsten
(168 tse27)
Coprinus satehtae Reid (1958: 430)
Coprinus rhombisporus Orton (1972: 145)

PILEUS 3-5 mm high unexpanded, narrowly paraboloid-conical,
white to whitish, sometimes tinted faintly brownish towards the
center, finely appressed floccose scaly towards the disc, becoming
pale mouse grey and vaguely striate when fertile, expanding to plane,
with edges becoming revolute. LAMELLAE ascending, nearly free, white,
becoming greyish-sepia, crowded, somewhat deliquescing with age.

STIPE 5-20 X 0.2-0.5 mm, subequal, white, nearly glabrous to
minutely pulverulent, with a flattened basal disc fringed with
tapering radiating mycelial strands.

PILEAR VEIL ELEMENTS: loosely interwoven, 2-4.5um diam.,
somewhat duplex, lower elements with a poorly developed coralloid
branching pattern, thin-walled, sometimes obscurely roughened, hyaline
or faintly brownish, tending to deliquesce early, upper or outer
elements normally well developed but sometimes nearly lacking, with a
well developed thorny to antler-like branching pattern, thick-walled,
having numerous elongated narrow branchlets which are solid or nearly
so, smooth to vaguely roughened; veil elements loosely attached to the
trama tissue which lacks a well developed pellis. PILEUS TRAMA:
hyphae radiating, 3-5yum diam., little inflated, thin-walled, hyaline,
smooth, with simple and clamped septa. LAMELLAR TRAMA: exceedingly
thin; 2-3 hyphae thick, loosely interwoven, and often septate, nearly
forming a pseudoparenchymatous layer, hyaline, thin-walled.
PLEUROCYSTIDIA: scattered, numerous, 70-75 X 14-20um, subcylindrical
to fusoid, thin-walled, hyaline. CHEILOCYSTIDIA: polymorphic, a
mixture of shorter pleurocystidia-like elements, swollen
brachybasidium-like elements and thin-walled veil-like elements
forming a mat over the lamellae edges. BASIDIA: dimorphic,

Figs. 1-12. Basidiospores. 1, C. psychromorbtdus DAOM 175229.

2, C. psychromorbidus DAOM 179229. 3, C. psychromorbidus DAOM 175232.
4, C. psychromorbidus ex Dearness DAOM. 5, C. psychromorbidus DAOM
189345. 6, C. psychromorbitdus DAOM 177600. 7, C. psychromorbtdus
DAOM 175226. 8, C. heritnktt Type at PRM. 9, C. pseudofrtestt Type at
PRM. 10, 11,C. mantholepra Type at Es »l2)) C.aphaeccsporue typerat Hs
All ca. 1,200 X mag.

B77

578

15-21 X 8.8-9.7um, 4-spored, subglobose with a short to long tapering
pedicel, spaced by swollen brachybasidia forming a rosette pattern.
BASIDIOSPORES: 7.5-9.3 X 5.5-7.9um, usually broadly ovate in face
view, varying to broadly elliptical, inequilateral in profile with a
less convex surface on the abaxial surface, thick-walled, smooth,
brown vinaceous in mass, Sienna in H,O, with a well developed apical
germ pore. STIPE HYPHAE: parallel, often inflated especially
internally, 3-15yum diam., hyaline, thin-walled, smooth.

HABITAT and SUBSTRATES: On leaf litter of Typha and Carex (Ont.) or
unspecified monocots (Que.) in wet fields or temporary pools.

COLLECTIONS EXAMINED: CANADA: Ontarto: Ottawa, June 28, 1979,
S.A.R. 3455 (DAOM 175138). Québee: Cantley, July 15, 1980, J. Ginns
(DAOM 177601). ENGLAND: Hertfordshire, King's Langley, Sept. 23,
1955, R.A. Saich (K, type for C. satchtae); Norfolk, Surlingham,
Wheatfen Broad, Aug. 3, 1968, P.D. Orton 3249 (E, type for

C. rhombisporus). SWEDEN: Vastergdtland, Gdéteborg, St. Anggarden,
Aug. 2 & 3, 1937, T. Nathorst-Windahl 658 (DAOM, Lundell & Nannfeldt
2928 as: C. jriesz2)..

An authentic Quélet collection of Coprinus frtestt could not be
located at Paris (PC). The concept adopted here is that of Pilat &
Svréek (1967), Lundell & Nannfeldt (1979), Kihner & Romagnesi (1953)
and Romagnesi (1937), i.e. a species with thick-walled, hyaline to
faintly yellowish, coralloid veil elements and broadly ovoid brownish
spores 7.5-9.3 X 5.5-7.9um.

The types for both Coprinus satchtae (figs. 16, 47) and
C. rhombtsporus (figs. 13, 45) do not differ significantly from other
collections of CC. friesit. There is a slight variation in the colour
of the veil elements, from hyaline to faintly yellowish, and in spore
Sizes but these are not unexpected considering the variations found in
the better documented case of C. psychromorbtdus. Nearly complete
sterility or a nearly complete arrestment of full veil development as
in DAOM 175138, make identifications difficult.

In Europe, Coprinus frtesit has been found on senescent
Trtfoltum, Lupinus, Chenopodtum, Arcttum, Robtnia, Carex and various
grasses (see references above, Orton & Watling 1979, and Lange 1915,
1939). It has also been reported from Morocco (Malengon & Bertault
1970) on Phragmttes and Juncus, and from Japan (Imazeki & Hongo 1965).
The report from Venezuela by Dennis (1961) is based on
CMOUSEROTYECSTUm devalue

*Coprinus kubickae Pilat & Svréek (1967, Aug.: 142) Figs. 67,68
= Coprinus amphtbius Anastasiou (1967, Nov.: 2213-4).

COLLECTION EXAMINED: CANADA: British Columbia: basidiocarps from a

Figs. 13-20. Basidiospores. 13, C. rhombtsporus Type E.

14, C. frtesit ex Lundell & Nannfeldt DAOM. 15, C. herbtvorus Type at
F. 16, C. satechiae Type at K. 17, C. suburttcicola Type at PRM.

18, C. urtteteola DAOM 175421. 19, C. brassicae Type at NYS.

20, C. melo, Type at G. ~All ca. J',200 X mag.

380

Figs. 21-24. Basidiospores. 21, C. maysotdtsporus DAOM 175231.
22, C.argenteus Type at K. 23, C. ttgrinellus Type at PC.
24, C. subtigrinellus Type at K. All ca. 1,200 X mag.

culture made from basidiocarps on wood, 4.5 mi. S of Clinton, by

Hwy. 97, July 24, 1964, L. Broome & N. Corfman (Anastasiou A 261 =
DAQM 128359, type C. amphtbtus).

An isolate of Coprinus kubtekae from turf grass in Saskatchewan
was recently described by Redhead & Smith (1981). A subsequent study
of the type of C. amphtbtus indicates that it is conspecific. Spore
size (fig. 67), colouration, reaction in H,SO, solution, and shape are
identical. Veil tissues (fig. 68) are similar although the hyphae are
less inflated on the average in the type of C. amphtbius. The habitat
differs as C. amphtbius was described on wood in alkaline lakes
whereas C. kubtekae is normally on herbaceous material. A wetland
habitat and possibly a basic substrate is common to all collections
known. This includes a collection from intertidal salt marsh plant
debris, on both woody twigs and grass from Willapa Bay, Washington,
U.S.A., Mar. 13, 1971, described by Van de Bogart (1975) under the

381

unpublished and provisional name, "Coprinus wtllapaensis" which
appears to be C. kubtickae.

*Coprinus maysotditsporus Redhead & Traquair, ECS eo ee 30.
sp. nov. 31-33 ,35/37,42-

Pileus 4.5-7.0 mm alt., conteus demum planus, candtidus,
squamatus; squamae brunneae, floccosae. Lamellae librae, candtdae
demum atrae, approximatae. Sttpes 6-21 mm long., 0.5-1.0 mm crassus,
@andidus, glaber vel floccoeus. Sporae 7-9 X 6-7.6 X 5.1-5. 3um,
atrae, pelluctdae, angulatae vel maysotdtformts, laeves; aptcewn poro
germinattvo; tuntea erassa. Pleurocysttdta 76-94 X 32-37 um,
fustformts, hyalina. Hyphae velt, ramosae; tunica hyalina,
tnerustata fusce.

Holotypus: tn solum prope Medtcagtnem sattvam, DAOM 175231.

PILEUS paraboloid-conical, 4.5-7 mm high when unexpanded, whitish,
with pale umber-coloured to pallid appressed floccose scales mostly
in the central region, becoming convex and greyish with age,
shallowly plicate-striate; edges eroding. LAMELLAE ascending, nearly
free, white becoming fuscous black, crowded, deliquescing with age.
STIPE 6-21 mm long, 0.5-1.0 mm wide, subequal, white, nearly glabrous
to slightly fibrillose-floccose, with a slightly flattened swollen
base.

PILEAR VEIL ELEMENTS: loosely interwoven, only slightly
inflated 5-10um diam., with a poorly developed coralloid branching
pattern, thin-walled, hyaline or brownish in segments where incrusted,
clamped, loosely attached to the trama tissue which lacks a well
developed pellis. PILEUS TRAMA: hyphae radiating, often inflated
with age, 6-18um diam., thin-walled, hyaline, constricted at septa,
smooth. LAMELLAR TRAMA: exceedingly thin, 2-3 hyphae thick; hyphae
loosely interwoven, 2.7-5.5ym diam., hyaline, thin-walled, clamped.
PLEUROCYSTIDIA: 79-94 X 31-37um, broadly ellipsoid to ventricose,
often forked or dimpled at the apex where embedded in the opposing
hymenium, thin-walled, hyaline. CHEILOCYSTIDIA: polymorphic ranging
from shorter versions of the pleurocystidia to brachybasidium-lLike,
mixed with or attached to hyaline veil-like elements forming a matted
cover on the lamellar edges. BASIDIA: dimorphic, 17-24 X 9.9-10.1lum,
4-spored, broadly ellipsoidal with a short to long narrow pedicel;
brachybasidia flattened, swollen, forming a rosette around each
basidium. BASIDIOSPORES: 7-9 X 6-7.6 X 5.1-5.3um, subangular and
broadly ovate to cordate or triangular in face view, slightly
flattened and elliptic to ovate in profile, similar in shape to the
kernels of Zea mays, thick-walled, fuscous black in mass, blackish but
not opaque under the microscope in H,0, readily dissolving in
concentrated H»,SO,, with a prominent apical germpore. STIPE HYPHAE:

' 5-30um diam., hyaline, with thin to pronounced smooth walls, more
inflated internally.

HABITAT and SUBSTRATES: On soil or artificial soil adjacent to
Medteago sativa taken from arid environments.

COLLECTIONS EXAMINED: CANADA: Alberta: nr. Brocket, Aug. 17, 1979,
J.A.T. 1334b (DAQM 175233); Lethbridge, freezing tests in greenhouse

382

from outdoor plots, April 11, 1979, J.A.T. 1287b, (DAOM 175231,
Typus). U.S.A.: Montana: Echo Lake, July 19, 1928, C.H. Kauffman
(DAOM 89696).

Coprinus maysotdtsporus is distinguished by its blackish
translucent characteristically-shaped spores and its coralloid,
thin-walled, brownish, incrusted veil elements. Its large spore size
is the only feature which readily distinguishes it from C. argenteus,
a species known from one collection in the British Isles. Kauffman's
collection was labelled "Coprinus (trtangulospora) n.sp. Kauff.
unpublished". No published record of this collection could be found.

Cultural data follows the taxonomic portion.

*Coprinus psychromorbtdus Redhead & Traquair Figs. 1-7,25,
sp. nov. 32,36,38-42.

Ptleus 4-9 mm alt., 5-12 mm latus, contcus demum planus, candidus,
squamatus; squamae brunneae, floccosae. Lamellae ltbrae candtdae
demum atrobrunneae, approximatae, deliquescentes. Sttpes 26-70 mm
long, 1-3 mm crassus, candidus, glaber vel floccosus. Sporae
(6.4-) 7, 2-8.8(-9.6) X (4.2) 5=6(4773) wm acroprivinege, alocves:
elltpsotdeae vel ovotdeae; aptceum poro germtnativo; tuntca crassa.
Pleurocysttdta 65-100 X 15-20um, fustformia, hyalina. Hyphae velt,
ramosae, flavobrunneae; tuntea crassa.

Holotypus: tn aptcalibus senescenttbus Medtcagints sattvae,
DAOM 179229.

HABITAT and SUBSTRATES: On necrotic or senescent parts of Medtcago
sativa L. (Alta.), Trtticum aestivum L. (Man.), Urttea dtotea L. ;
(Man.), Elymus ptpert Bowden (B.C.), Agrostis stolontfera L. (Sask.),
Dianthus sp. (Ont.) and other herbaceous plants usually in arid
macro-environments.

COLLECTIONS EXAMINED: CANADA: Alberta: nr. Brocket, Aug. 17, 1979,
J.A.T. 1334a (DAOM 175232); Lethbridge, field plots, Sept. 20, 1979,
J.A.T. 1341 (DAOM 175228), freezing test materials removed from field
plots, April 11, 1979, J.A.T. 1286 (DAOM 175226), J.A.T. 1287 (DAQM
175230);,-April 17,, 197975 ALT.) L291 (DAM 175227) eo Ae ee 2
[Holotypus] (DAOM 179229), also culture from J.A.T. 1292 on horse
manure (DAOM 177647). British Columbia: 2 km. N. of Vernon,

Sept. 25, 1980, S.A.R. 4054 (DAOM 177600). Manttoba: Winnipeg,

Aug. 3, 1937, coll. unknown (DAOM 189344, 189345). Saskatehewan:
Saskatoon, in greenhouse material from outdoor plots, Mar. 12, 1981,
J. Drew Smith (DAOM 179672). Ontario: London area, Sept. 8, 1926,
"Emma" [Dearness] (DAOM).

A more detailed description of the basidiomes and cultures based
on Alberta collections was presented by Traquair (1980).

Coprinus psychromorbtdus is characterized by having thick-walled

Figs. 25-31. Basidiomes. 25, C. psychromorbtdus DAOM 177600.
26, 27, C. urtictcola DAOM 175421. 28, 30, 31, C. maysotdtsporus
DAOM 175231. 29, C. frtestt DAOM 175138.

384

Figs. 32-33. Mycelia in petri plates. 32, C. maysotdtsporus DAOM
175237 on upper halves versus C. psychromorbidus DAOM 175227 on lower
halves. 33, C. maysotdtsporus on PDA (left) and MA (right) 3 wks.

brownish antler-like veil elements and brown, thick-walled ovoid to
ellipsoid spores, mostly 7.2-8.8 X 5-6um. ‘The size, shape and
intensity of the melanization of the basidiospores is quite variable.
In the Ontario collection the spores were nearly all ellipsoidal and
relatively weakly pigmented, the Manitoba collections have spores with
prominently domed adaxial surfaces and in some cases a second germ
pore on this raised portion, the British Columbian collection had many
misshaped spores, was partially sterile and in general had elongated
Ovoid spores. The collections from Alberta showed a range in shapes
from broadly ovoid to ellipsoidal and in degrees of pigmentation.
Cultures from all of the Alberta collections interbred (Traquair
1980). The variation in spore characteristics probably indicates

Fig. 34. C. friestt, basidiospores, basidia, pleurocystidia, veil
elements from pileus, DAOM 177601. Scale = 15,m.

386

that the species has an unstable gene pool resulting in a high degree
of plasticity.

The earliest known collection of Coprinus psychromorbtdus was
made by Mrs. Emma Dearness, the wife of Dr. John Dearness. The
collection's label reads, "Coprinus brassicae... Emma D. found it on
stems of garden pink 8th Sept. 1926," with no additional locality data
given. This collection is considered to be from London, Ontario, John
Dearness's hometown (Tamblyn 1955) as Dearness collections at DAOM
(Parmelee 1978) made at more distant localities are so labelled
whereas collections made in the London area usually are not. In
addition, J. Dearness, and presumably his wife, had been home in
southern Ontario in September 1926 as is evident from his
publications (Dearness 1972a,b) and other miscellaneous collections.

Coprinus psychromorbtdus was next collected by W.F. Hanna (1939)
who reported it as C. urttctcola. He had earlier cited it as
C. phaeosporus (Bisby et al. 1938). Hanna's material on wheat and on
the nettle collected in July 1937, are not available for study. Two
collections labelled Coprinus brasstcae (a name Hanna considered
synonymous with C. urttctcola) made on herbaceous stems collected
behind the Dominion Rust Research Laboratory, Winnipeg, in Aug. 1937
by an anonymous collector are (. psychromorbidus. Also, Hanna's
(1939) description, line drawings and photographs of the collections
on wheat indicate that he had C. psychromorbidus and not C. urttetcola
or any other species. Thus Hanna was the first to describe the
species in culture and recognize that it might be of importance to
agriculture although he could not demonstrate any pathogenicity.

Coprinus psychromorbtdus was finally linked to snow mold damage
when dikaryotic-monokaryotic matings were made between sterile
isolates of the causal agent and single spore isolates from
basidiocarps of the Coprinus from Alberta (Traquair 1980, 1981). The
presence of one pathogenic species in section Herbtcolae raises the
possibilities that other species are pathogens. Coprinus kubtckae,
was found to be associated with superficial fairy rings on turf in
Saskatchewan and is possibly the causal agent (Redhead and Smith 1981).
Alternatively, the turf damage may have been caused by
C. psychromorbtdus, or C. kubtekae in combination with
C. psychromorbidus as the last species has also been isolated from the
affected turf (see specimens examined).

Coprinus psychromorbidus appears to be closely allied to the
European C. phaeosporus Karsten. Its spores are smaller, fall within
a narrower range, and the shape is more constant probably as a result
of the small size. Coprinus psychromorbtdus and C. phaeosporus
probably represent a vicarious species pair derived from a common
ancestral circumpolar population. The main difference is spore size
and that being highly variable for the former although not to the
extent of overlapping. In the same way that C. psychromorbtdus is a
large-spored equivalent of C. phaeosporus, C. maysotdisporus is a
large-spored equivalent of C. argenteus. These last two species
otherwise being nearly identical in habit, habitat and morphology,
also can be considered as a vicarious species pair. In the cases of
Coprinus frtesit, C. urttctcola and C. kubtekae which are all found on
both continents, more stable gene pools must be involved.

Rien

OO era

a Cece
ane

ae
OOO
eee

Fig. 35. C. maysoidisporus, basidiospores, basidia, veil elements
from pileus, DAOM 175231. Fig. 36. C. psychromorbidus basidiospores,
DAOM 179229. Scale = 15um.

388

Other closely allied species are C. herbtvorus, differing by
angular spores, and C. cinchonensis (q.v.), differing by large
spores.

*Coprinus urttctcola (Berk. & Br.) Buller Figs. 17-19,26,
(1917: 485) 27 ,56-62,69.
= Agaricus urttetcola ee & Broome

(1861: 376) ut "urtteaecota" (Art. 73.8, Stafleu Ce Cleo Oye
= Psathyra urtictecola (Berk. & Br.) eoevaa (1887 et 0 Je

Coprinus brasstcae Peck (1890: 64)

Coprinus melo Favre (1948: 215)

Coprinus suburticicola Pilat & Svréek (1967: 140)

to ou ol

PILEUS 3-4 mm high unexpanded, paraboloid-conical to nearly
hemispherical, white, nearly opaque, minutely pulverulent, becoming
plane with revolute torn edges, greyish sepia with age. LAMELLAE
nearly free, crowded, white, becoming greyish sepia with age,
deliquescing slightly. STIPE 6-11 mm long, 0.1-0.4 mm wide,
subequal, fistulose, white, minutely floccose to nearly glabrous,
arising from a basal disc with a sparse fringe of tapering mycelial
strands.

PILEAR VEIL ELEMENTS: loosely interwoven, with a poorly to
moderately developed coralloid branching patter, little inflated,
5-10um diam., thin-walled, hyaline, sometimes with refractive
hardened drop-like incrustations, simple-septate; loosely attached to
the trama tissue which lacks a well developed pellis. PILEUS TRAMA:
exceedingly thin; hyphae radiating, thin-walled, hyaline, 2-5um
diam., little inflated, simple-septate. LAMELLAR TRAMA: exceedingly
thin, 2-3 hyphae thick; hyphae interwoven and often septate forming a
nearly pseudoparenchymatous tissue, 2-5um diam., hyaline, thin-walled.
PLEUROCYSTIDIA: abundant, scattered, 65-88 X 16-24um, fusoid to
ventricose, thin-walled hyaline. CHEILOCYSTIDIA: polymorphic,
varying from shorter pleurocystidia-like elements to swollen
brachybasidium-like elements and veil-like elements binding the
lamellae edges together. BASIDIA: dimorphic 13-22 X 6.5-7.0um,
4—spored, sSubglobose to utriform, with a short to long narrow pedicel,
simple-septate. BASIDIOSPORES: 5.5-8 X 3.9-5.lum, narrowly to
broadly ellipsoidal to obscurely ovoid, sometimes approaching
subglobose in shorter spores, slightly inequilaterally flattened on
the adaxial surface, smooth, thin to very slightly thick-walled, pale
brown under the microscope, sepia in mass, with a prominent transverse
to tangential apical germ pore. STIPE: hyphae parallel, more
inflated intermally, 14-15um diam., thin-walled, hyaline, smooth.

HABITAT and SUBSTRATES: On Carex (Que.) or grass leaves in wet open
fields.

COLLECTIONS EXAMINED: CANADA: Québec: Cantley, June 24, 1979,
S.A.R. 3490 & J.H. Ginns (DAOM 175421), June 29, 1980, S.A.R. 3518 &
J.H. Ginns (DAOM 177602). CZECHOSLOVAKIA: Horni Slovénice pr.
Lommice nad LuZnici, Bohemiae, July 14, 1962, M. Svréek & J. Kubictka
(PRM #567917, type for C. suburttetcola). ENGLAND: Kew, July 24,
1957, D.A. Reid (DAOM 66599, ex K). ITALY: Athesis fl. (= Adige
River), Sept. 1900, G. Bresadola (NY). SWITZERLAND: Jura vaudo's

PAG. Oo L«
Scale =

C. maysotdisporus, pleurocystidia, DAOM 175231.
15um.

218.9

SHA,

vallée de Joux, prés du Sentier, Riére la C6te, Aug. 28, 1939, J.
Favre (G, type of C. melo). U.S.A.: Wew York, Albany Co., Menands,
Aug. C.H. Peck (NYS, type for C. brasstcae); Bronx, June 15, 1911,
W.A. Murrill (NY).

The type of Agaricus urticicola, the basionym of Coprinus
urtictcola, was unavailable for study because of its fragmented
nature. Orton (Dennis, Orton & Hora 1960: 222; Orton 1960: 410)
illustrated and described the spores from the type as ellipsoidal,

6-8 X 4.5-5.5yum, with an apical germ pore. This is in rough agreement
with Cooke's (1886) illustration from the type collection showing
ellipsoidal to obscurely ovoid spores, 7 X 4ym. The combination of
spores of this type and a pure white pileus for a European Coprinus

in section Herbicolae indicates that the name should be applied to

the species as accepted here.

The types of Coprinus brasstcae (Figs. 19,58), C. melo (Figs.
20,57) and C. suburtictcola (Figs. 17,59) all have hyaline,
thin-walled coralloid veil elements and ellipsoid to obscurely ovoid
brownish spores with a prominent apical germ pore. Coprinus brassicae
is clearly synonymous with C. urtictcola. Many of the spores of
C. suburtictcola are larger than typical C. urtictcola but the ranges
overlap too much to allow maintaining them as separate taxa. The
position of the germ pore is also too variable. Coprinus melo differs
mainly in the gross appearance of the pileus, having prominent
rib-like folds suggesting a type of melon. The microscopic features
are identical to typical C. urticteola. The type of C. melo would
seem to be a growth form of C. urttcetcola possibly effected by the
high altitude environment.

Coprinus urtictcola has been reported on Brasstca oleraceae L.
and Zea mays L. in the U.S.A. (Peck 1980 & Murrill 1912, as
C. brassteae) and on Urtteca dtotea L., Carex, Juncus, branches of
Picea abtes Karst. and various grasses including Phalarts arundinacea
L. and Agropyron species in Europe (Pilat & Svréek 1967).

The report of Coprinus urticicola from Manitoba by Hanna (1939)
was based on material of C. psychromorbidus (q.v.). Pegler's (1977)
report from east Africa is based on (. paleotroptcus (q.v.) and the
Venezuelan collection reported by Dennis (1961) is C. neotroptcus
(deVies

EXTRALIMITAL SPECIES
*Coprinus argenteus Orton (1972: 139) Figs. 22;oL

COLLECTION EXAMINED: ENGLAND: Surrey, Mickleham, Juniper Hill,
June 22, 1956, C. Kemball (K, type).

This is the European small spored equivalent of
C. maysotdtsporus. It has blackish but translucent spores readily
dissolving in concentrated H2SO,. The spores are identical in shape
to C. maysotdtsporus, but are smaller, 5.5-6.5 X 5.0-5.2 X 4.5-4.8um.
The veil elements are moderately to poorly coralloid, essentially
thin-walled and hyaline with patches of hyphae bearing a thin
brownish incrustation.

591

Fig. 38. C. psychromorbidus veil elements, cheilocystidia (center),
pleurocystidia, basidia, DAQM 179229. Scale = 15um.

The habitat is on grass in chalkland grasslands.

*Coprinus austrofrtesit Redhead & Pegler, sp. nov. PAG -moos

Differt a Coprtno friesto sports ovotdets 9.6-11.8 X 7.2-8 xX
7-9um.
Holotypus: tn graminis, Iulto 11, 1958, Los Palos Grandes (Caracas)
Edo, Miranda, Venezuela, leg. R.W.G. Dennis 1495 (K).

Coprinus austrofrtesit was reported from Venezuela as C. friestt
by Dennis (1961: 120). Dennis evidently noted a difference between
his collection and European materials as he suspected it would be
segregated at the species level in time. It differs from C. frtestt
by the much larger spores. In other respects it resembles C. friestit.
This is an important difference when compared to the consistency
shown between European and North American collections of C. frtestt.

*Coprinus burkit Smith in Smith & Hesler (1946: 178).

The type of this species was not studied. This species is
essentially C. phaeosporus (q.v.) but with hyaline veil elements
which have thinner walls towards the extremes of the branches than in
C. phaeosporus. The habitat is on fern petioles in Alabama.

*Coprinus ctnehonensts Murrill (1918: 85) Fig2, 547%

COLLECTION EXAMINED: JAMAICA: Cinchona, 4500-5200 ft. level,
Dec.25-Jan. 8, 1908-9, W.A. & E.L. Murrill 579 (NY, type).

Very little is known about this species whose name is based on a
Single basidiome. The fact that it grew on a log may indicate that it
is a peripheral species to the section Herbtcolae, however, our and
Smith's (1948) type studies showed that the microscopic features are
very similar to the fungus described here as C. psychromorbtdus.
Murrill's species differs by having larger spores, 9-ll X 5-6 X
6-7.5um and by the lignicolous habitat. The veil elements are
slightly paler yellow than most C. psychromorbtdus veil elements.

*Coprinus herbtvorus Singer (1973: 70) _ Figs. 15,49.

COLLECTION EXAMINED: ARGENTINA: Jujuy, Lagunas de Yala, Feb. 9,
1965, R. Singer T3929 (F1015235, type).

Singer (l.c.) gave a detailed description of this species. The
angularity of the spores, the most characteristic feature of this
species, varies from prominent to obscure. Veil elements with a
prominent linear axis from which branched elements arise also
characterize C. herbivorus.

BITS,

Figs. 39-47. Basidiospores. 39, C. psychromorbidus DAOM 189345.
40, C. psychromorbidus DAOM 177600. 41, C. psychromorbtdus ex
Dearness DAOM. 42, C. psychromorbidus DAOM 175239.

43, C. maysoidisporus DAOM 89696. 44, C. friestt DAOM ex Lundell &
Nannfeldt. 45, C. rhombisporus Type at E. 46, C. friestt DAOM
175188. 47, C. satchiae Type at K. Scale = 15ym.

394

*Coprinus herinkii Pilat & Svréek (1967: 137) Figs. 8,66.

COLLECTION EXAMINED: CZECHOSLOVAKIA: southern Bohemia, Cernohéj nr.
Vodnany, Aug. 12, 1938, J. Herink (PRM 499770, type).

Only one collection of this species is known but it seems
distinct enough to warrant recognition. The small subglobose spores
with apical wrinkles and patches, if found to be a consistent
character, will readily distinguish it from all other taxa.

*Coprinus neotroptcus Redhead & Pegler, sp. nov. EIQ eos

Differt a Coprino urtictcolo sports elltpsotdets vel ovoidets
8-9,5 X 6.5-7. dum.
Holotypus: in Bambusets, Iulto, 4, 1958, El Avila, Dts. Federal,
Venezuela, leg. R.W.G. Dennis 1136 (K).

Coprinus neotroptcus was reported from Venezuela as C. brassicae
by Dennis (1961: 118-119). As with C. austrofriesit (q.v.) he
suspected that the collection would be segregated from the northern
taxon. Coprinus neotropicus differs from C. urtietcola by having
larger spores. It is, in other respects, very similar to the latter
species.

*Coprinus paleotroptcus Redhead & Pegler, sp. nov. Eigemoze

Differt a Coprino urttctcola sports ovotdets, 6.1-8 X 4.5-5 X
5.8-6.7um, tuntcts, crassts.
Holotypus: «in ltgno, Martto 25, 1968, Nyanza Prov., Kericho reg.,
Kigumu fl., Kenya, leg. D. Pegler 234 (K). Paratypus: Pegler 368.

Coprinus paleotropteus is well described and illustrated as
C. urttetcola by Pegler (1977: 397-399). It differs from
C. urtitetcola by having more prominently ovoid, thicker-walled spores
and by the woody substrate.

*Coprinus phaeosporus Karsten (1881: 9) Figs: 9-12,63-65.
= Coprinopsts phaeosporus (Karsten) Karsten
(1881b:7 27)

Coprinus pseudofriesii Pilat & Svréek (1967: 140-141)
Coprinus xantholepts Orton (1972: 150)

fo ofl

COLLECTIONS EXAMINED: CZECHOSLOVAKIA: central Bohemia,
Praha-Divoka, July 22, 1941, J.A. Herink 332/41 (PRM 626346, type of
C. pseudofriestt). FINDLAND: southern Tavastia, Tammela, Mustiala,
Aug. 5, 1880, P.A. Karsten 1602 (H, lectotype of C. phaeosporus).
SCOTLAND: Aberdeenshire, Loch Skene, Aug. 19, 1964, P.D. Orton (E,
type of C. xantholepts).

The data for the collection which was designated as lectotype
and illustrated by Horak (1968) agrees well with that of the
protologue (Karsten 1881) and a subsequent description (Karsten
188lc), i.e. Mustiala, in terra stercoratis, 5 Aug. 1880. The type

Figs. 48-55. Basidiospores. 48, C. subtigrinellus Type at K.
49, C. herbivorus Type at F. 50, C. tigrinellus Type at PC.
51, C. argenteus Type at K. 52, C. paleotroptcus Type at K.
53, C. neotroptcus Type at K. 54, C. cinchonensis Type at NY.
55, C. austrofrtesit Type at Kk. Scale = 15m.

consists of a number of basidiomes, some cespitose on grassy debris.
As noted by Horak (1968) and Pilat & SvrGek (1967), the spores
(Figs. 12,65) from the extant basidiomes are smaller, 5.5-7 X 5.5-6um,
and shaped differently from what Karsten had observed, i.e. 9-15 X
5-9um. This discrepancy is best explained, as suggested by Pilat &
Svrcek (l1.c.), by assuming that Karsten had a mixed collection and
what now remains, represents only one of the species represented in
the original collection. Thus the species as lectotypified is one
characterized by having veil elements with a well developed
antler-like branching pattern, thick, golden to hyaline refractive
walls, and brownish thick-walled spores which are broadly ovate in
face view and slightly compressed, inequilateral and with a nearly
lateral apiculus in profile.

The name Coprinus xantholepts (Figs. 10,11,64) is based ona
collection with essentially identical features to C. phaeosporus as is
C. pseudofrtestt (Figs. 9,63). There is a slight difference in the
degree of pigmentation of the spores in the three types but not enough
to segregate them into different taxa, especially when one considers
the variation observed in compatible strains of the allied
C. psychromorbtdus. The report of C. phaeosporus from Manitoba
(Bisby et al. 1938) is based on C. psychromorbidus (q.v.).

*Coprinus subttgrinellus Dennis (1961: 122-123) Figs. 24,48.

COLLECTION EXAMINED: VENEZUELA: Caracas, Botanic Garden, June 30,
1958, R.W.G. Dennis 1124 (K, type).

As indicated by the name, C. subtriginellus is allied to
C. tigrinellus, both having veil elements which are brownish,
essentially thin walled and coralloid.

Coprinus subtigrinellus differs from C. tigrinellus in having
smaller spores, 5.7-7 X 4.5-6um vs. 8-10 X 6.8-7.lum. Otherwise they
are very similar although the former is known from South America and
the latter from Europe.

*Coprinus ttgrinellus Boudier (1885: 283) Pigs ./#237507

COLLECTIONS EXAMINED: FRANCE: Montmorency, July 1884, E. Boudier
(Lectotype marked "Icones N. 409") plus two additional collections
from 1884 and 1890 (PC).

The veil elements of C. ttgrinellus are not as rounded as
illustrated by Boudier (1905-1910) but more like that illustrated for
C. subttgrinellus by Dennis (1961). The walls are relatively thin
and vary from nearly hyaline in some segments to quite darkly
melanized in others. They do not become thickened and refractive or
golden as in C. phaeosporus or C. psychromorbtdus and they are not
roughened by incrustations to the extent that the veil is in
C. argenteus or C. maysotdtsporus.

3O7

Figs. 56-67. Basidiospores. 56, C. urttctecola DAOM 66599.

57, C. melo Type at G. 58, C. brasstcae Type at NYS.

59, C. suburttcicola Type at PRM. 60, C. urtictcola DAOM 175421.
61, C. urticicola NY ex Murrill. 62, C. urtictcola NY ex Bresadola.
63, C. pseudofriestt Type at PRM. 64, C. santholepts Type at E.

65, C. phaeosporus Type at H. 66, C. herinkit Type at PFM.

67, C. amphibius Type at DAOM. 68, Veil elements of C. amphtbtus.
Scale = 15um.

398

PROBLEMATIC NAMES
*Coprinus frtestt var. mtcrospora J. Lange (1915: 42).

No type collection exists for this name and insufficient data was
given in the original description to determine its taxonomic position.

*Coprinus mtcrospermus J. Lange (1939: 112), nom. invalid.

This is a nomen nudum, nomen provisiorum and no voucher specimens
exist. It is probably the same as C. friesit var. mtcrospora but no
connection was stated.

*Copritnus pallidtsporus J. Lange (1939: 112), nom invalid.

This is a nomen nudum, nomen provisiorum and no voucher specimens
exist.

*Coprinus platypus Berkeley in Cooke (1888: 234; 1886: pl. 687B,
[675B]).

The type of this species is deposited at Kew. It was described
from a collection on palm leaves in conservatories in England and was
considered as an alien species. Microscopic details from the type
recorded by Dr. D.N. Pegler at Kew (pers. comm.) indicate that the
spores are 7.5-9.5 X 6-7.5um and the veil elements are branched in a
manner similar to that for C. friesti. The elements have thick
hyaline walls and are 1.5-4.5um diam. Additional observations made by
one of us (S.A.R.) on a lamellar fragment indicate the presence of”
pleurocystidia, an abundance of simple-septa, 4-spored basidia lacking
clamps, and brownish slightly thickened walls on the spores. The
shape of the spores is subglobose to broadly ellipsoidal.

This species keys out to C. frtestt and is probably a synonym of
that species but the total lack of clamp connections, and the more
rounded spore shape without obvious ovoid forms as seen in all other
collections of C. friestt raise a degree of doubt. Additional
collections of C. friesit showing more of an intergradation with
C. platypus should be studied before the latter name is permanently

placed in synonymy.

*Coprinus phaeosporus var. solitarius J. Lange (1939: 111),
nom. invalid.

This is a nomen nudum and no voucher specimen exists. Pilat &
Svrcek (1967) proposed the combination Coprinus solitartus (Lange)
Pil. & Svr. based on Lange's variety. However, the combination is
based on an invalid name and even though the latin description in
Pilat & Svrcek's key couplet might suffice for validation, no type
was designated as was required. The name thus remains invalid.

S99

Fig. 69. C. urtictcola, veil elements from pileus, cheilocystidium,
pleurocystidia, basidiospores, basidia, DAOM 177602. Scale = 15,m.

400

*Coprinus picosporus Locquin (1955: 13), C. pusto Locquin (1955: 16)
and C. romagnest Locquin (1955: 14).

These three species no doubt belong in Coprinus section
Herbtcolae but insufficient details were published and no types were
designated to allow comparisons. In addition, Reid (1958) has noted
that the spore sizes given in the text do not correspond with the
measurements calculated from data from the illustrations. He
suggested that if the magnification stated for the illustrations of
spores were changed to 1000X from 2000X there would be consistent
correlations. If this were done all three would be taxa distinct from
others in section Herbicolae. However, if the magnifications given
for the illustrations of spores is correct, i.e. 2000X then the spores
would be 1/2 the size reported in the text. Coprinus ptcosporus
would have spores 4.5-6 X 3.5-4.2um. Based on spore shape and the
thick-walled pigmented coralloid veil elements, C. pitcosporus would
become a synonym of C. phaeosporus. Coprinus romagnest would have
spores 8-9.5 X 6.5-8.5ym and presuming that the veil elements were
thick-walled (data not given), then C. romagnest would be synonymous
with C. friestt. Coprinus pusto would have spores 7-8 X 4.5-5um.
Presuming that Locquin's collection had a poorly developed veil,

C. pusto would be synonymous with C. urtictcola.

Until the characters to be associated with Locquin's three names
can be determined, the use of the names is problematic.
Lectotypifiable material may be present in Dr. Locquin's herbarium
but this was unavailable during its transfer to the Laboratoire de
Micropaléontologie, Paris (Locquin, pers. conm.).

CULTURAL STUDIES

The following information on cultures of C. maysotdisporus
should be compared with that given for C. psychromorbidus (see
Traquair 1980). The isolate (DAOM 175231) was obtained by streaking
a spore suspension, obtained from a powdered dried basidiome, on
potato dextrose agar. The procedures follow Nobles (1965).

Macroscopic description:

Growth moderate to rapid at 22°C, covering 9-cm petriplates in
2-3 wk (8.0 mm/da), with optimal temperature 25°C; advancing zone
even to uneven, thin, appressed; aerial mycelium white, appressed to
felt-like and somewhat patchy and plumose after 3-4 wk (Fig. 33),
occasionally producing yellowish to pale brownish patches of densely
matted hyphae after 4-5 wk; surface and submerged mycelium producing
white, globose to irregular hyphal knots, 1-3 mm wide, which develop
into brownish-black, smooth sclerotia at 6 wk; reverse unchanged in
color; odor not distinctive; no evidence of fruiting on malt extract
agar or potato dextrose agar.

Tests for extracellular oxidases with gallic acid were positive
with no subsequent mycelial growth and were positive, changing color
within 2 min with alcoholic gum guaiacum.

Tests for hydrogen cyanide with picric acid solution and PDA as
growth medium (Lebeau and Hawn 1963) were negative after 45 da.

401

Microscopic description:

Hyphae hyaline, thin-walled, noninflated and with clamp
connections unless otherwise noted; hyphae of the advancing zone
2.0-3.5um wide, occasionally branched; hyphae on the agar surface
2.0-4.5(-5.0) um wide, frequently branched and loosely interwoven after
2 wk, becoming more frequently branched and interwoven after 3-4 wk;
submerged hyphae 2.5-4.0(-10.0) um wide, frequently branched, gnarled,
contorted, with globose, elliptical, or irregular swellings and having
thin to slightly thickened hyaline to yellowish refractile walls.
Hyphal knots and sclerotia compactly interwoven, internal hyphae
(medulla) of sclerotia pseudoparenchymatous, 4.0-12.0um wide, hyaline,
thin-walled, inflated, containing granular or oily contents; outer
hyphae (rind) of sclerotia plectenchymatous, 4.0-8.0(-12.0) um wide,
yellowish to brown-walled, slightly thickened to thick-walled,
cylindrical to irregularly inflated.

Sexuality studies:

Spores from basidiocarps:-of collection DAOM 175231 germinated in
4-12 h on potato dextrose agar. Hyohae of the monokaryotic, primary
mycelium lacked clamp connections but otherwise resembled the

dikaryotic, secondary mycelium.

Coprinus maysotdtsporus like Coprinus psychromorbtidus, is
heterothallic and is probably bipolar. This pattern is indicated by
the complete compatibility of 12 single spore isolates with the
dikaryotic isolate in di-mon pairings. Such a result is consistent
with findings for other bipolar basidiomycetes (Raper 1966).
Tetrapolar species usually have 50% compatibility in intraspecific,
intracollection di-mon matings.

‘The di-mon matings between C. maysotdisporus and
C. psychromorbtdus did not result in the dikaryotization of the
monokaryotic mycelia, thus supporting the view that two species
should be recognized. Monokaryotic mycelium of each species was not
dikaryotized by the other Coprinus species but was dikaryotized by
dikaryotic isolates of the same species. Moreover, colonies of
different species failed to merge. The different colonies remained
distinct and a confrontation or barrage zone of mounded hyphae
developed at the interface between them. ‘

Discussion on the cultural characteristics:

Coprinus maysotdtsporus differs from C. psychromorbidus in
colony morphology and optimal growth temperature. The former produces
appressed mats whereas C. psychromorbidus produces woolly or cottony
mats. Unlike the latter Coprinus, the former has a higher temperature
optimum (i.e. 25°C) and grows very slowly if at all, at temperatures
below 10°C. Furthermore, C. maysoidisporus does not produce HCN in
culture as does C. psychromorbidus (Traquair 1980).

The hyphae of the two species are similar. Although
allocyst-like and antler-like branches characteristic of the hyphae
of C. psychromorbidus were not observed in C. maysotdtsporus, it would
be extremely difficult to distinguish cultures of the two species on
the basis of hyphal anatomy.

402

Whether or not C. maysotdtsporus is pathogenic is not known at
present.

ACKNOWLEDGEMENTS

We thank the following curators and mycologists for their
invaluable assistance by the loan of type and other materials: Drs.
J. Haines (NYS), B. Spooner, D. Pegler and D. Reid (K), C. Rogerson
(NY), D. Henderson and R. Watling (E), H. Harmaja (H), M. Svréek
(PRM), O. Monthoux (G), S. Jovet and J. Mouchacca (PC), and P. Ponce
de Leon (F). We also wish to thank Dr. J. Ginns for collecting fresh
materials and commenting on the manuscript and Dr. M. Locquin for
correspondence concerning his collections. J. Drew Smith kindly
reviewed the manuscript, as did R. Watling, and provided the
Saskatchewan collections of C. psychromorbtdus. Dr. D. Pegler
provided invaluable assistance at Kew. Mrs. M. Meredith typed the
final manuscript.

LITERATURE CITED

Anastasiou, C.J. 1967. ‘Two species of Coprinus from alkali lakes.
Canoe POb en a 5c e2 i= 222.
Berkeley, M.J. & C.E. Broome. 1861. Notices of British Fungi.
Ann. Mag. Nat. Hist., ser. 3, 7: 373-382.
Bisby, G.R., A.H.R. Buller, J. Dearness, W.P. Fraser, R.C. Russell &
H.T. Gussow. 1938. The fungi of Manitoba and Saskatchewan.
N2R.C. CG, Ottawa.
Boudier, E. 1885. Description de quelques espéces nouvelles de
Champignons basidiosporés. Bull. Soc. Bot. Fr. 32: 282-285.
. 1905-1910. Icones Mycologicae. Lib. Sci. Natur., Paris.
Broadfoot, W.C. 1936. Experiments on the chemical control of
snow-mould of turf in Alberta. Sci. Agric. 16: 615-618.
Broadfoot, W.C. & M.W. Cormack. 1941. A low-temperature
basidiomycete causing early spring killing of grasses and
legumes in Alberta. Phytopathology 31: 1058-1059.
Buller, A.H.R. 1917. Some critical remarks on the generic position
of Psathyra urttcaecola Berk. et Broome, Coprinus pltcattlis
Fr., and Psathyrella disseminata. Trans. Brit. Mycol. Soc.
5: 482-489.
Cooke, M.C. 1886. Illustrations of British Fungi (Hymenomycetes).
No. XXXIX and No. XLIII. Williams and Norgate. London.
. 1888. Handbook of British Fungi. [published with
separate pagination in Grevillea]. Agaricini No. 513-949.
Grevillea 16(80): ut 145-256.
Dearness, J. 1927a. Amantta poisoning. Mycologia 19: 93-96.
. 1927b. Fleshy fungi. Mycologia 19: 228-229.
Dennis, R.W.G. 1961. Fungi Venezuelani. IV., Kew Bull.
E532 67-156.
Dennis, R.W.G., P.D. Orton & F.B. Hora. 1960. New checklist of
British agarics and boleti. Trans. Brit. Mycol. Soc. 43
(suppl. )
Favre, J. 1948. Les associations fongiques des hauts-marais
jurassiens et de quelques régions voisines. Beit.
Kryptogamenflora Schw. 10(3): 1-228, I-IV pls.

403

Hanna, W.F. 1939. Coprinus urttcaecola on stems of Marquis wheat.
Mycologia 31: 250-257.

Horak, E. 1968. Synopsis generum Agaricalium (Die Gattungstypen der
Agaricales). Beit. Kryptogamenflora Schw. 13: 1-741.

Imazeki, R. & T. Hongo. 1965. Coloured illustrations of Fungi of
Japan. Vol. II. Hoikusha Publ. Co., Osaka.

Karsten, P.A. 188la. Symbolae ad mycologiam Fennicam. VIII.
Meddel. af Soc. pro Fauna et Flora Fennica 6: 7-13.

- 1881b. Hymenomycetes Fennica. Acta Soc. pro Fauna et
flora Fennica. II, 1: 1-40.

- 188lc. Symbolae ad mycologiam Fennicam. IX. Meddel.
af Soc. pro Fauna et flora Fennica 9: 39-56.

Kuhner, R. & H. Romagnesi. 1953. Flore analytique des champignons
supérieurs. Masson et Cie, Paris.

Lange, J.E. 1915. Studies in the agarics of Denmark. Pt. II.
Amantta, Leptota, Coprinus. Dansk. Bot. Ark. 2(3): 1-53.

. 1939. Flora agaricina Danica. IV. Danish Bot. Soc.,
Copenhagen.

Lebeau, J.B. 1969. Diseases affecting forage production in western
Canada .0ub. 125-1338. sel Kor. Nielson, ed... Proc., Can.
Forage Crops Symp., Western Cooperative Fertilizers Ltd.,
Calgary.

Lebeau, J.B. & E.J. Hawn. 1963. Formation of hydrogen cyanide by
the mycelial stage of a fairy ring fungus. Phytopathol.

5339 1395-1396.

Locquin, M. 1955. Recherches sur les Coprins. Bull. Soc. Mycol.
ry eee Om LO.

Lundell, S. & J.A. Nannfeldt. 1979. Fungi Exsiccati Suecici.
Praesertim Upsalienses. Fasc. LVII-LX (Nr. 2801-3000).
Reklam & Kalalogtryck, Uppsala.

Malengon, G. & R. Bertault. 1970. Flore des Champignons superieurs
du Maroc. I. Trav. Inst. scient. chérif. et Faculté des
Sci., Rabat., ser. bot. biol. végét. 32: 1-604.

Porrill, W.A. 1912... Illustrations of. Fungi.- X. Mycologia 4: 1-6.

. 1918. The Agaricaceae of tropical North America -
Waals issyerelherests, AUOR weyy toler

Nobles, M.K. 1965. Identification of cultures of wood-inhabiting
Hymenomycetes. Can. J. Bot. 43: 1097-1139.

Orton, P.D. 1960. New checklist of British agarics and boleti.
Part III. Notes on genera and species in the list. Trans.
Bi Wey COOL OO Ee LOU SSO ¢

. 1972. Notes on British agarics: IV. Notes Royal Bot.
Gard?) sloinburdh pe 322 3b=L50.

Orton, P.D. & R. Watling. 1979. British fungus flora. Agarics and
Boleti. 2. Coprinaceae Part 1: Coprinus. Roy. Bot. Gard.
Edinburgh, Edinburgh.

Parmelee, J.A. 1978. The Dearness mycological collection.
Mycologia 70: 509-526.

Peck, C.H. 1890. Annual report of the state botanist (1889). From
the 43d report of the New York State Museum. Bot. ed.,

p. 1-48, 4 pls:

Pegler, D.N. 1977. A preliminary agaric flora of east Africa. Kew.
Bull ye ocd ty ser.nvi.

Pilat, A. & M. Svréek. 1967. Revtsio specierum secttonts Herbtcolae
Pil. et Svr. generis Coprinus (Pers. ex) S.F. Gray.

Ceska Mykol. 21: 136-145.

404

Quélet, M.L. 1872. Les champignons du Jura et des vosges. Mém. Soc.
Emul. Montbéliard, ser. II, 5: (43-332).
Raper, J.R. 1966. Genetics of sexuality in higher fungi. Ronald
Press Co., New York. 283 p.
Redhead, S.A. & J.D. Smith. 1981. A North American isolate of
Coprinus kubtekae associated with a superficial fairy ring.
Gan. Js Bot. 59: 410-4147 ae
Reid, D.A. 1958. New or interesting records of British
hymenomycetes II. Trans. Brit. Mycol. Soc. 41: 419-445.
Romagnesi, H. 1937. Florule mycologique des Bois de la Grange et de
L'Etoile. Rev. Mycol, 2: 243-255.
- 1951. Etude de quelques Coprins. Rev. Mycol.
16: 108-128.
Saccardo, P.A. 1887. Sylloge Hymenomycetum I. Agaricineae.
Patavia.
Singer, R. 1973. Diagnoses fungorum novorum Agaricalium III.
Beih. Sydowia 7: 1-106.
Smith, A.H. 1948. Studies in the dark-spored agarics. Mycologia
40: 669-707.
Smith, A.H. & L.R. Hesler. 1946. New and unusual dark-spored
agarics from North America. J. Elisha Mitchell Sci. Soc.
62: 177-200.
Stafleu, et al. 1978. International code of botanical nomenclature.
Regnum Vegetabile 97: 1-457.
Tamblyn, W.F. 1955. John Dearness. Mycologia 47: 909-915.
Traquair, J.A. 1980. Conspecificity of an unidentified snow mold
basidiomycete and a Coprinus species in the section
Herbicolae. Can. J. Plant Path; @: 105-115.
1981. Discovery of a sporulating state for the LTB
snow mold. Can. Agric. 26: 18-20.
Van de Bogart, F. 1975. The genus Coprtnus in Washington and
_adjacent western states. Ph.D. thesis, Univ. Wash.

NOTE ADDED WHEN IN PRESS

A few additional collections from Edinburgh (E) were examined
Which are of interest. Coprinus herbtvorus is now ‘known from
Australia: Sydney, April 6, 1974, R. Watling Herb. No. 10892, on
grass. Two paratypes of C. rhombtsporus were examined. The earliest,
Orton No. 962, July 18, 1956, consists of what appears to be a mixed
collection of C. friesit with slightly abnormal spores (some slightly
rhomboid) and C. phaeosporus. The second collection, Orton No. 4134,
Aug. 30; 197 ASC. nyrveses.

MYCOTAXON

WiOd) wid [eNO 2 3 pp...405-406 July-September 1981

SCLEROTINIA BRESADOLAE RICK, A TAXONOMIC SYNONYM
OF CIBORINIA CANDOLLEANA (LEV.) WHETZEL

LINDA M. KOHN

Department of Botany, Clemson University
Clemson, SC 29631

Since type or authentic material of Sclerotinia bresado-
lae Rick (Inoperculate Discomycetes) was unavailable
during the author's previous studies of the genus Scle-
BOULa,) NO, COnCclustVve: disposition. sol, thes Species ~could
Dewmade and the epithet “was. listed in “A “monographic
revision of the genus Sclerotinia’ (Kohn, 1979), as "im-

perfectly known." Recently the holotype became avail-
able for study and disposition of the Species as a taxo-
nomic synonym of Ciborinia candolleana (Lév.) Whetzel

can now be made.

Rick (1900) described the species from a_ collection
Pride wine soprino, aloo0, . ‘One rotting. ‘buds sci, Quercus,
many of which were infected by @a gall-forming wasp,
Deyoteras» terminalis.iHe “distinguished the new. Species
maomumoclerotinia, candolleana Lév., <which occurs on
leaves of Quercus, primarily on the basis of what he
Fell twas Valunique: substrate, ~ buds ‘transformed into
Pols seine dddition,. hes cited the longer stipessand@ithe
hairs borne on apothecia and stipes as distinctive.

This species clearly fits within the accepted circumscrip—
Mor Ole cl porinia Whetzel. “Though ™=hypertrophtedy due” fo
insect infection, the host is foliar. The ectal excipulum
er the ‘apothecium is composed of Jarge-celled” textura
globulosa. The loaf-shaped sclerotia incorporate host

xylem vessels distinguishable by their spirally thick-

ened walls; the medullary cells are dorsiventrally dif-
ferentiated with the upper zone composed of textura glo-
bulosa with gelatinous walls, and the lower zone com-
posed of textura oblita. No conidial anamorph has been
observed for this species.

This species also fits within modern circumscriptions of

406

Ciborinia candolleana. Stipe length is within the range
for that species as described by Batra (1960) and, ac-
cording, tom Rickas scomments™ ono.) Diesadolae, 515 ane
doubt due to a thick layer of litter covering the sclero-

tia of this particular population. Batra mentions "rhi-
zoidal tufts present'' on the stipe, and stout, rhizoidal

hyphae, (ane Mpresen inva ta ieauDaSemol sl iema pote! uin Ole
ten connecting directly to the sclerotium, in Rick's ma-—
terral. ~NOw cthen, Behdarniung sumatha la cou immacCOUn a aOm
Ricks, “description Stipites o.com Vil OSO,EViGESUSEmdlsCuUln
fére i lanucginoso ss wanOWws sev 10Cll UY mee x CCD Umm OCmELULL omen
fine jhyphae,= originating either sirom oermindtingmspomes

or from infection of apothecia by another fungus, seengl

in some older apothecia examined.

Because particular weight has been placed on host spe-
cCificity, in) delimitation) sol fspeciesmin wy Ciborimian tien cca
CUrGeENCes Ol § Ogu Dresadolac mon mOUCTCUSMEIS MUI DOGLAD mann
considered as meristematic leaf tissue, buds and galls
are “probably, “not ssignificantly sidifferent = (rome themicaves
of Quercus, saithouc hpricks cao pecimenmda ppcausm OmuecpEC.
Sent) GaN graves occurrences OhmsO we candollca lia mCi ccen
galls.

SPECIMEN EXAMINED: Rick, Majo 1889, "in gemmis Quer-
cus, transformatis. intectione | Dreyoteralis sterminakisow melols
land. [S, ex herb. Bresadola; slide in CUP 59836).

ACKNOWLEDGEMENT

Appreciation’ 15 sexpressed tome) es ochumachiel mun iverciiy
of Oslo, for making this specimen available for study.

DORR RAT OR Ea Griieb

BAGUREy gael a Rie 1960. The species of Ciborinia patho-
genic to Salix, Magnolia, and Quercus. Amer. |.
Bot. 47: 819-827.

KOHN, L. M. 1979. A. monographic revision of the ge-—
nus Sclerotinia. Mycotaxon 9: 365-444.

Ru Ree |. 1900 Eine neue, Sclerotinia—Art. Oesterr.
inven WAGE ASUS Tar

MYCOTAXON

Vol. XIII, No. 2, pp. 407-411 July-September 1981

HALYSIOMYCES, A NEW DEMATIACEOUS GENUS
FROM ARIZONA'S SONORAN DESERT

EMORY "G. SIMMONS

Department of Botany, University of Massachusetts
Amherst, Massachusetts 01003

Clumps and chains of dark-walled, nonsporulating
fungus cells are extremely common inhabitants of practical-
ly any surface that offers a bit of nutrient. Substrate
examples include discarded cellulosic products, painted
surfaces, almost every scrap of plant debris overwintered
in temperate climates, and the surfaces of living leaves
and stems, especially in the tropics but also in temperate
climates when aphid secretions and similar nutrients afford
support.

Identification of such nonsporulating growths rarely
is possible. But the simple expedient of holding a sample
in a damp chamber for a day or so frequently induces sporu-
lation and thus enhances the possibility of identification.
Species of Cladosporium, Aureobasidium, and Alternaria com-
monly develop from such materials in temperate climates;
other less easily characterized genera also may be found.

Discussed herewith is a species that, judging from its
Characters in culture, probably will be seen in nature pri-
Mariya heateall)isasechainsvand clumps: of dark, xounded,
thick-walled cells. It grows readily as an isolate on weak
potato—carrot agar, hay decoction agar, and 202 V-8 juice
pear "(henreartervas PCA,{Hay, V-83 see Stevens 1974)%0 It
produces conidia abundantly and of a septate, catenulate
kind that is believed to be remarkable for this heteroge-
neous group of dematiaceous molds. Inspiration to discuss
this species has derived largely from staff publications of
the Centraalbureau voor Schimmelcultures (Hermanides-Nijhof
1977; Hoog and Hermanides-Nijhof 1977) and of the University
of Alberta Mold Herbarium and Culture Collection (Sigler,
Tsuneda and Carmichael 1981), which treat several super-
ficially similar genera.

408

HALYSIOMYCES Simmons, gen. nov. Hyphomycetes
[Gr.: halysion + mykes = chain-fungus]

Hyphae atrobrunneae, torulosae, ramosae. Colonia atra
ut radii discreti vel. fila radiata arcte adpressa formata;
plerumque in substrato agaro submersa, sed ad centrum ali-
quantum elevata, convoluta, cerebriformis; aspectu nitido
sed nec uda nec mucosa. Hyphae aeriae infrequentes. Coni-
dia ex cellulis hypharum holoblastica, atra, transverse
septata, singularia vel distaliter catenata.

Typus: Halysiomyces saxatilis Simmons, sp. nov.

Described from axenic culture. Hyphae dematiaceous,
torulose, branching. Colony black, developing as discrete
radii (on PCA and Hay) or as closely appressed radial ele-
ments; mostly submerged in agar substrate, but raised, con-
voluted, cerebriform in center; appearing shiny on V-8 but
not wet or mucoid. Aerial hyphae rare. Conidia holoblastic
from hyphal cells, dark, transversely septate, distally
catenate.

Halysiomyces saxatilis Simmons, sp. nov.
[L.: saxatilis = found among rocks]

Ex culturis in agaro PCA descripta. Coloniae ut hyphae
atrobrunneae, torulosae, ramosae, submersae, valde radiatae
formatae. Centrum coloniae convolutum, aliquantum elevatum,
hyphis aeriis paucis. Cellulae hypharum laeves, usque ad
6-l6um longae x 16-20um latae ampliatae. Conidia ex cellu-
lis hypharum radialium et ramulorum lateralium holoblastica,
allantoideoclavata, minute aspera, brunnea, 1-3 transverse
septata, circa 25 x 4um, et solitaria et 2-4-catenulata
acropeta.

Origo typi: ex aere deserti Sonora prope Phoenix,
Arizona, U.S.A. /im.) Febr.| 1980,avJd. TT. Staley et Paeraimer
lecto.

Typus: partes ex Simmons 33-129 desiccatae et in BPI,
DAOM, IMI, NY conservandae.

The isolate is described from cultures grown concur-
rently on PCA, Hay and V-8 juice agars. Radial growth is
about 10mm in 7 da at 15-22 °C in diurnal light/dark cycles.

Fig. 1. Halysiomyces saxatilis. Thick-walled radial and
branch hyphae with solitary and catenate conidia produced
as submerged elements in PCA.

410

Radiating torulose hyphae produced in PCA are distinct
and somewhat sinuous; concentric rings of growth correlate
with the light-dark regimen; the colony center is slightly
raised above the agar surface and is closely convoluted
under pressure of progressive septation, enlargement, and
hardening of the toruloid elements. The macroscopic ap-
pearance on Hay agar is similar to that on PCA except that
the radial pattern of growth is less striking than are the
medusoid radii produced in PCA. The radial elements pro-
duced on V-8 agar are appressed so closely that the colony
appears opaque; there is a larger, more obviously convolut-
ed central area of development than is found on PCA; and
the agar surface has a dull sheen which, on inspection at
low magnification, proves to be light reflected from a net-
work of jetblack, closely septate torulose hyphae resem-
bling millipede integuments or doliiform beads in chains of
indeterminate length. Colonies are dry, not mucoid or wet.

Linear hyphal growth is of brown tubular elements that
gradually become septate and torulose-swollen at more or
less regular intervals; most resultant cells are 10-20um
broad and shorter than wide; a longitudinal septum is laid
down in relatively few hyphal cells. Lateral branches of
one to many swollen brown cells are numerous throughout the
length of the radial hyphae, but they do not obscure the
predominantly radial appearance of most colonies. Otherwise
undifferentiated cells of radial hyphae and of lateral
branches may produce conidia, sometimes only one or two per
conidiogenous cell but at other times clusters of 6-10.

Primary conidia are sessile, pale, minutely rough, and
slightly enlarged toward the apex; they darken, produce a
median transverse septum, and become inequilateral, usually
with an abrupt bend near the distal end; at the 1l-septum
stage they commonly are 25 x 4um, becoming only slightly,
if at all, larger when second and third transverse septa
are formed. Conidia may develop a subapical or truly later-
al extension 2-3yum wide x 1-3um long which functions as the
site of production of a single holoblastic conidium. Chains
of 2-4 or more conidia are produced in this manner; they are
easily visible as they are held in position submerged in the
agar substrate. Conidium ontogeny appears to be holoblastic
in all cases. Conidium release appears to be by wall frac-
ture, a small remnant of conidiogenous cell wall sometimes
being visible alongside the bottom contour of a dislodged
conidium.

Production of distinctive conidia submerged in agar
substrates, as with this fungus, is a striking phenomenon;
but, in my experience, it is not unique to this fungus, in

All

that some strains of Stemphylium and Curvularia have been
observed to sporulate abundantly and with typical conidium
morphology under such conditions.

Information from isolator J. T. Staley is that this
fungus (typified from an air isolate) also has been found
in culture plates prepared from swabbings of a material
referred to as "rock varnish," which is found on rock sur-
faces in the cactus-creosote bush region of the Sonoran
desert near Phoenix, Arizona, U.S.A.

-oO0o0-

Margaret E. Barr Bigelow and I had several fruitful
discussions on the systematic placement of this isolate.
J. W. Carmichael, after reviewing a draft of the manuscript,
has commented on unique aspects of the species.

= (Q) =
LITERATURE CITED

Hermanides-Nijhof, E. J. 1977. Aureobasidium and allied
genera. CBS Studies in Mycology, no. 15, p. 141-177.

Hoog, G. S. de, and E. J. Hermanides-Nijhof. 1977. Survey
of black yeasts and allied Hyphomycetes. CBS Studies
eM COlLOo VO Lo mn Oo2 2 27

Sigler, L., A. Tsuneda and J. W. Carmichael. 1981. Phaeo-
theca and Phaeosclera, two new genera of dematiaceous
hyphomycetes and a redescription of Sarcinomyces Lind-
ner. Mycotaxon 12(2): 449-467.

Stevens, R. B. (ed.) 1974. Mycology Guidebook. Seattle.
TASB Gok

MYCOTAXON

Vol. XLIL, ‘No, 2, .pp.. 412-415 July-September 1981

OSORIOMYCES, A NEW GENUS OF THE LABOULBENIALES
FROM TAIWAN

KATSUYUKI TERADA

Botantcal Instttute, faculty of Sctence,
Htroshima Untverstty, Htroshtma 780, Japan

SUMMARY

A new species of the Laboulbeniales parasitic on
Osortus formosae Bernhauer (Coleoptera, Staphylinidae) is
described as Osortomyces rhtzophorus. Osortomyces is a
new genus having an affinity with Scelophoromyces; it is
characterized by a thallus consisting of a primary axis of
superposed cells that terminates in a cluster of appendages;
this axis bears sessile perithecia, as well as secondary
axes on which short antheridial appendages and perithecia
are borne laterally. Rhizoidlike appendages grow downward
from a cluster of small cells just above the basal cell of
the thallus. The outer walls of the perithecia consist of
four vertical rows of four cells each. Although spermatia
could not be detected, structures having the appearance of
simple antheridia were observed.

During my investigation of the Laboulbeniales of Taiwan,
a fungus was found that bore a resemblance to Scelophoro-
myces osortanus Thaxter, which had been collected on
Osortus in Argentina (Thaxter, 1912, 1931). However, 'the
perithecia of the Taiwanese thalli had very short stalk
cells, giving the appearance of being sessile, and the
thalli bore secondary axes. Consequently, the following
new genus is being described. Both fungus and host speci-
mens are deposited in the Herbarium of Hiroshima University,
Hiroshima, Japan (HIRO).

Osortomyces Terada, gen. nov.

Thallus ex axe primario. et axibus secundariis constans.
Axis primarius ad apicem appendicem terminalem et secus

413

latera appendices laterales et prope basin rhizoidea gerens.
Axes secundarii secus-latera appendices laterales gerentes. ~
Cellulae receptaculi praeter cellulas prope basin uniseri-
ales. Perithecia sessilia in axe primario et axibus sec-
undariis lateraliter prodientia. Parietes exteriores
perithecii ex seriebus 4-cellularibus quattuor constantes.
Antheridia simplicia in appendicibus lateralibus prodientia.

Thallus consisting of primary and secondary axes.
Primary axis bearing primary appendage at its apex, lateral
appendages along its sides, and rhizoidlike appendages near
its base. Secondary axes bearing lateral appendages.

Cells of receptacle (except for cells in the suprabasal por-
tion of the primary axis) arranged in a uniseriate row.
Sessile perithecia produced laterally on both the primary
and secondary axes. Outer walls of perithecia consisting
of four vertical rows of four cells each. Simple anther-
idia produced on lateral appendages.

Type species: Osortomyces rhtzophorus Terada, on
Osoritus formosae Bernhauer from Taiwan.

Osortomyces rhtzophorus Terada, sp. nov. (Figs. 1-18)

Thallus hyalinus vel leviter luteolus, 320-540 um
longus, in receptaculi 2-5 perithecia gerens. Axis) p ri—
Hmarius ex 25-35 cellulis superpositis oblongis.constans.
Axes secundarii ex 8-13 cellulis superpositis qblongis con-
stantes. Appendix terminalis ca. 50 um longa, plerumque
ramosa, gelatinosa, sparse septata, basi septo denigrato
constricta. Appendices laterales 26-52 um longae, simpli-
ces vel varie ramosae, gelatinosae, sparse septatae, sursum
extensae. Rhizoidea 180-350 um longa, simplicia, rigidula,
clare septata, deorsum extensa. Antheridiasca. 9, [Lim longa,
evanescentia, gelatinosa, solitaria vel fasciculata.
Perithecia 60-79 x 24-36 um, ovata, plerumque extrinsecus
curvata, apice truncata.

Thallus hyaline or slightly yellowish, 320-540 um long
(from foot to constricted, dark septum at base of terminal
appendage), bearing 2-5 perithecia on receptacle. Primary
axis consisting of 25-35 superposed, oblong cells; cells
just above cell I undergoing several longitudinal and trans-
verse divisions to form group of cells. Secondary axes
shorter and thinner than primary axis, on which they are
irregularly arranged, each secondary axis consisting of 8-
13 superposed, oblong cells. Terminal appendage ca. 50 um

414

Figs. 1-18. Osortomyces rhtzophorus. 1. Sporeling with
second and third cells divided longitudinally. 2. Young
thallus, showing receptacle with two rhizoidlike appendages
(ra), a lateral appendage (la), and a terminal cluster of
appendages. Arrow indicates blackened septum. Dee UppeL
portion of primary axis}, showing a cluster of appendages;
only terminal one bears blackened septum at its base.

4. Upper portion of primary axis, showing a cluster of ap—
pendages; terminal one also bears blackened septum at its
base. 5. Upper portion of primary axis, showing appendages
produced on both sides of each receptacle cell. Left arrow
indicates a subdichotomously branched appendage. Right
arrows indicate antheridia with or without a one-celled
stalk. 6. Upper portion of secondary axis, showing the
initiation of an appendage below and a triangular, subses-
sile cell above bearing a one-celled antheridial branch with
three apical antheridia. 7. Short trichotomous appendage.
Arrow indicates triangular basal cell. 8. Secondary axis,
showing a short, simple appendage. Arrow indicates pri-
mary axis. 9-11. Very early stages of perithecial devel-
opment. Explanation is in text. 12. Young peritheciun,
showing the female organ, which consists of three cells -

a terminal trichogyne (tr), a medianttrichophoric cell (e.),
and a carpogenic cell (f). 13. Immature perithecium at
3-tier stage, showing a trichophoric cell (e'), a superior
supporting cell (ss), an ascogenous cell (ac), a secondary
inferior supporting cell (ist), and an inferior supporting
cell (is). 14. Fully mature perithecium on primary axis,
with appressed stalk cell (VI). Lines indicate septa in
Vertical row, Ole Outera walt ceLis, 15. Mature perithecium
in which the fourth tier has divided in one row. Line in-
dicates dividing point. 16. Slightly immature perithecium
on secondary axis; primary axis (arrow) bends to left.

17. Thallus with two well-developed perithecia and two sec-
ondary axes (arrows). 18. Thallus with two mature perithe-
cia and one young perithecium; five rhizoidlike appendages
extend from suprabasal portion of primary axis; gelatinous
lateral appendages arise from upper portion of primary axis;
secondary axis (arrow) has broken. (Figs. 1-16: scale B;
Figs. 17-18: scale Ay)

415

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416

long, usually branched, gelatinous, sparsely septate, con-
stricted by blackish septum. at its base. Lateral append-
ages 26-52 um long, simple or variously branched, gelati-
nous, extending upward, arising from triangular, subsessile
cells formed at one or both upper corners of receptacle
cells, these appendages sometimes producing one or more
antheridia terminally. Simple rhizoidlike appendages 180-
350 um long, somewhat rigid, extending downward from more
or less cubical cells in suprabasal cluster of receptacle
cells. Antheridia ca. 9 um long, evanescent, gelatinous,
solitary or clustered, produced directly upon somewhat tri-
angular basal cell or borne on short, one-celled branch.
Perithecia 60-79 x 24-36 um, ovate, usually curved outward
and truncate at its apex.

Holotype: TAIWAN. Nantou Hsien, Chitou, at ca. 1300 m
alt., No.w)/83,) May 293) 19V7. Keeleradallec..20n elytrago.
Osorius formosae Bernhauer (Coleoptera, Staphylinidae).

Paratypes: TAIWAN. Nantou Hsien, Chitou, Nos. 782, 7/84,
786-790, data as for the holotype; Nantou Hsien, Tsuifeng,
Nos. 791, 792, July 25% 1972) (Kew leradagles. .oOngvaktous
parts of body of Osortus formosae Bernhauer.

Osortus fotmosae occurs in mountainous regions of Tai-
wan, where it lives in rotten wood. It is often found
with Prtochtrus stlvestrit Bernhauer, host of Monotcomycés
leptochtrt Thaxter (Terada, unpublished). Fichtatully
mature specimens of Osortomyces and more than ten immature
ones were collected from thirteen specimens of Osortus
formosae. Collection of additional specimens would be
necessary for a thorough study of thallus development.

After spore germination, a series of horizontal divi-
sions takes place that results in the formation of a uni-
seriate thallus having a darkened, constricted septum near
its apex, which may break off early (Fig. 1). Vertical
divisions soon occur in the lower cells of the thallus,
resuiting in the formation of a biseriate lower. receptacle
CEL mel The number of cells in the thallus increase so
that in the thallus shown in Fig. 2, there are eight super-
posed cells separating the constricted, dark septum from
the cluster of cells just above the basal cell. Rhizoid-
like appendages (ra) extend outward just above the base of
the thallus, whereas gelatinous, erect appendages (la) arise
laterally at a higher level. The primary axis of the thal-
lus terminates in a cluster of gelatinous appendages (Figs.
2-4) (it was not possible to determine the exact position
of the spore septum, which separates the primary appendage

417

from the receptacle). Byamaturitysetive on six. simple,
long, rhizoidlike appendages are present that extend down-
ward from the suprabasal group of cells in the primary axis
cs / sald). Triangular, subsessile cells are formed
on one or both upper corners of the upper cells of the re-
ceptacle. Each of these cells produces a short, gelati-
nous branch. Secondary axes that arise from the upper
part of the primary axis (two are indicated in Fig. 17) may
deteriorate later (Fig. 18). The positions of the perithe-
cia are variable; however, the lowermost one usually occurs
on the suprabasal portion of the primary axis (Fig. 18).

In the material examined, one specimen bore gelatinous,
unicellular, elongate structures directly on triangular,
subsessile cells (Fig. 5) or in clusters on a stalk cell
produced by a subsessile cell (Figs. 5, 6). Although sper-
matia could not be detected, these structures are regarded
as simple antheridia because they are very narrow apically.
Consequently, Osortomyces belongs to the Laboulbeniaceae.
The antheridia seem to cease their function in a short time;
possibly, they are converted into sterile appendages (Fig.
Dea GLOwraleloeht shies ou) AlO)ie

During perithecial ontogeny in Osortomyces rhtzophorus,
a single cell (a') that arises laterally from the recepta-
cle (Fig. 9) soon divides obliquely into an upper (d) and
aelower celli'(c) (Fig. 10). The lower cell (c) divides
PiCowanwupper cel lacG )mandhaw lower, Celle (ce) (FhTo bn),
which undergo further divisions and finally form the peri-
ehecial iwalls. The upper cell (d) produces the female sex-
ual organ, which initially consists of three superposed
cells - a lower carpogenic cell (f), a median trichophoric
Pou (em) ew andeal terminals trichogyne, (tr) (Fig. 12). Only
one— or two-celled trichogynes were observed. By the time
there are three tiers of outer wall cells, an ascogenous
cell (ac) and three supporting cells (ss, ist, is) are pres-
ent below the trichophoric cell (e") within the perithecium
(higt 3). This pattern of perithecial development is the
Same as that described in detail by Thaxter (1896) for
Stigmatomyces baert (Knoch) Peyritsch and other taxa.

By the time perithecial wall consists of four tiers of
cells, asci have been formed (Fig. 16). The second tier
of cells in the fully mature perithecia is 3-4 times taller
than the lowest tier (Fig. 14). Sometimes the apical out-
er wall cell on the inner side of the perithecium (toward
Ene thallus) axis) divides (Fig. 15). The stalk cells and
the basal cells are strongly appressed to the receptacle so

that the perithecia are sessile in appearance (Figs. 14,
15).

418

The genus Osortomyces cannot be allied with any genus
of the Laboulbeniales except Scelophoromyces Thaxter (1912;
in 1931, Thaxter altered the spelling of this genus to
SkeLophoromyces, but the earlier spelling should be used).
Osortomyces resembles Scelophoromyces in the following ways:
1) the primary axis consists of a variable number of super-
posed cells; 2) rhizoidlike appendages arise at right and
left from the suprabasal portion of the primary axis; 3)
lateral appendages extend upward from one or both upper cor-
ners of receptacle cells. On the other hand, remarkable
differences exist between the two genera: 1) the perithecia
of Osortomyces are normally sessile and lateral, whereas in
Scelophoromyces, they are long-stalked and never arise di-
rectly from any cell of the primary axis; 2) the primary
axis of Osortomyces is highly divided in the suprabasal por-
tion, whereas in Scelophoromyces, the suprabasal portion is
less divided or undivided; 3) lateral appendages of Osorto-
myces are gelatinous and the protoplasts of their short,
subsessile basal cells taper almost to a point apically,
whereas in Scelophoromyces, they are rigid like the rhizoid-
like appendages and the protoplasts of their basal cells
are broad at the apex.

ACKNOWLEDGMENTS

I wish to thank Dr. I. I. Tavares, University of Cali-
fornia, Berkeley, for reviewing the manuscript as well as
for helpful suggestions and Mr. Y. Shibata, Tokyo, for
identification of the ,jhost insects.

LITERATURE CITED

Thaxter, (R.as1890; Contribution towards a monograph of the
Laboulbeniaceae. Mem. (Amer soAcad Ss 7AntsaSc in lZ sao
L298 Pls. I-XXVIL.

Pe tie Pap New or critical Laboulbeniales from the
Argentine. ProcsvAmern. (Acad. Arts @SCint 46 2553-22,
LOS Ts. Contribution towards a monograph of the
Laboulbeniaceae. Payiaas Mem. Amer. Acad. Arts Sci.
16: ,1-435% Bboy lak.

MYCOTAXON

Moles WiyeNO.e25 pp. 419-427 July-September 1981

THE GENUS CODINAEA. THREE NEW SPECIES FROM THE AMERICAS

ADRIANNA D. HEWINGS

Department of Plant Pathology, University of Illinois,
Urbana, IL 61801

J-L. CRANE

Illinois Natural History Survey and the Departments of
Botany and Plant Pathology, University of Illinois, Urbana,
IL 61801

Three new species of Codinaea Maire are described and
illustrated from the U.S.A. and Venezuela. A key modified
from Hughes and Kendrick (3) to include those species
described since 1968 is also included.

Codinaea illinoensis Hewings & Crane SD.enOVs ieee.

A and B.

Coloniae effuse, pallide brunneae, demum
fusco-brunnescentes. Mycelium in substrato immersum, e
hyphis crasse tunicatis, ramosis, septatis, subhyalinis vel
pallide brunneis compositum. Setae nullae. Conidiophori
macronemati, mononemati, simplices, recti vel paulum
arcuati, septati, hyalini vel pallide brunnei, tunicis
crassis, levibus induti, singillatim e hyphis vel cellulis
nodulosis orti. Cellulae conidiogenae monophialidicae,
integratae, terminales, paulum lageniformes, apicem versus
tenuiter, prope basim crasse tunicatae, 30-37 x 5.6-6.7 um.
Collarula conspicua, infundibuliformia, expansa, marginem
versus tenuiter tunicata, ad basim crasse tunicata,
refringentia, 2.3-3.3 x 3.3-4.4 um. Phialoconidia hyalina,
continua vel nonnumquam uniseptata, paulum arcuata,
symmetrica, ad apicem attenuata in finem obtuse rotundatum
setulam rudimentarium apiculatum exhibentem, ad basim
attenuata, cicatrice obtusa notata, 14.4-18.9 x 3.3- 4.4
um. Typus: ILLS 42169.

420

Colonies effuse, pale brown becoming dark brown.
Mycelium immersed see the substrate, composed oD
thick-walled, branched, septate, subhyaline to pale brown
hyphae. Setae absent. Conidiophores macronematous,

Fig. 1. Codinaea illinoensis, A. Conidiophores;

eae a ee

B. Mature Conidia.

A21

mononematous, simple, straight or slightly curved, septate,
hyaline to light brown, thick-walled, smooth, arising
Singly from the hyphae or nodulose cells. Conidiogenous
cells monophialidic, integrated, terminal, slightly
lageniform, thin-walled at apex, thick-walled at base,
30-37 x 5.6-6.7 um. Collarettes conspicuous, funnel-
shaped, flaring, distally thin-walled, thick-walled and
refringent at base, 2.3-3.3 x 3.3-4.4 um. Phialoconidia
hyaline, unicellular or occasionally 1-septate, slightly
curved, symmetrical, tapering to a bluntly rounded distal
end bearing a rudimentary apiculate setula, basal end
tapering with a blunt scar.

Holotype: On decayed wood, cypress swamp, N.W. of
Vienna, Johnson County, Illinois, 9 July 1969, J.L. Crane
98-69, ILLS 42169.

Codinaea lunulospora Hewings & Crane sp. nov. Fig.2, A-F

Coloniae effusae, pallide demum obscure brunneae.
Mycelium in substrato immersum, e hyphis crasse tunicatis,
ramosis, septatis, subhyalinis vel pallide brunneis
compositum. Setae nullae. Conidiophori e basibus sculptis
orti, macronemati, mononemati, simplices, recti, septati,
obscure brunnei, crasse tunicati, leves, ad phialidis basim
paulum attenuati. Cellulae conidiogenae phialidicae,
integratae, terminales, lageniformes, ad apicem tenuiter
tunicatae, hyalinae, crasse tunicatae, subhyalinae vel
pallido brunneae ad basim. Collarula conspicua,
infudibuliformis vel aliquantulum cylindrica, apicem versus
tenuiter tunicata, sympodialiter semel vel bis instaurata,
3.6-4.5 x 1.8-2.7 um, prope basim ad 1.4-1.8 um attenuata.
Phialoconidia hyalina, continua, lunata vel sigmoidea,
symmetrica, 8.8-12.0 x 0.8-1.0 um. Holotypus: Dumont-VE
4619 (NY). Isotypus: (VEN).

Colonies effuse, light brown becoming dark brown.
Mycelium immersed in the substrate, composed Of
thick-walled, branched, septate, subhyaline to light brown
hyphae. setae absent. Conidiophores arising from
sculptured bases, macronematous, mononematous, Simple,
Straight, septate, dark brown, thick-walled, smooth,
tapered slightly at base of phialide. Conidiogenous cells
Pphialidic, integrated, terminal, lageniform, thin-walled
and hyaline at apex, thick-walled, subhyaline or light
brown at base. Collarettes conspicuous, funnel-shaped to
somewhat cylindrical, distally thin-walled, developing
sympodially with 1-2 successive proliferations, 3.6-4.5 x

A422

’

Codinaea lunulospora, A-E. Conidiophores

Fig.

Mature Conidia.

423

1.8-2.7 pm narrowing to 1.4-1.8 um at base. Phialoconidia
hyaline, unicellular, crescent-shaped or Sigmoid,
symmetrical, 8.8-12.0 x 0.8-1.0 um.

Holotype: On unidentified herbaceous stem. Trail
between Manacal and Los Pocitos, N.W. of Irapa, Edo Sucre,
Venezuela, 10 July 1972. K.P. Dumont et al., Dumont
VE-4619 (NY), Isotype: (VEN).

Codinaea matsushimae Hewings & Crane sp. nov. Fig. 3, A-D

Coloniae effusae, pallide brunneae, demum obscure
brunneae. Mycelium in substrato immersum, e hyphis crasse
tunicatis, ramosis, septatis, subhyalinis vel pallide
brunneis compositum. Setae singillatim vel gregatim ortae,
rectae, crasse tunicatae, leves, ad vicies septatae, ad
basim obscure brunneae, ad apicem obtuse rotundatum pallide
brunneae vel subhyalinae, usque 340 um longae, e basi 6.7
um lata ad apicem 4.4 um lata attenuatae. Conidiophori e

cellularum nodo setarum socio orti, macronemati,
mononemati, simplices, recti vel paulum arcuati, septati,
hyalini vel pallide brunnei, crasse tunicate, leves.

Cellulae conidiogenae phialidicae, integratae, terminales,
hyalinae, paulum lageniformes, ad apicem tenuiter, ad basim
crasse tunicatae, 14-27 x 3.0-6.0 um. Collarula conspicua,
evanescens, ad marginem tenuiter, ad basim abrupte crasse
tunicata, synpodialis, semel, bis, vel ter instaurata,
2e0=3-0. x 2.0-4.0 im. Phialoconidia setulis ornata,
hyalina, ter ver raro semel vel bis septata, paulum
arcuata, symmetrica, ad apicem obtuse rotundatum et basim
cicatrice notatum attenuata, 20-27 x 3.3-5.4 um. Setulae
alia ad apicem, alia juxtam cicatricem basalem affixae,
5.5-7.8 um longa. Typus: ILLS 42170.

Colonies effuse, light brown becoming dark brown.
Mycelium immersed in the substrate, composed of
thick-walled, branched, septate, subhyaline to light brown
hyphae. setae arising singly or in groups, erect,
thick-walled, smooth, up to 20-septate, dark brown at base,
pale brown to subhyaline at bluntly rounded apex, up to 340
um long, tapering from 6.7 um wide at base to 4.4 um at
apex. Conidiophores arising singly from a knob of cells
associated with setae, macronematous, mononematous, simple,
Straight to slightly curved, septate, hyaline to light
brown, thick-walled, smooth. Conidiogenous cells
phialidic, integrated, terminal, hyaline, slightly
lageniform thin-walled at apex, thick-walled at base, 14-27
x 3.0-6.0 um. Collarettes conspicuous becoming evanescent,

424

Codinaea matsushimae, A-C. Conidiophores and

Ficw 3,

D. Mature Conidia.

setae;

425

distally thin-walled, abruptly thick-walled at base,
Ssympodial with 1-3 proliferations, 2.0-3.0 x 2.0-4.0 um.
Phialoconidia with setulae, hyaline, 3-rarely 1 or
e-septate, slightly curved, symmetrical, tapering toa
bluntly rounded distal end and a proximal basal sear, 20-27

x 3.3-5.4 um. Setulae attached at distal end and to one
side of basal sear, 5.5-7.8 um long.
Type: On exocarp of Carya, Benson’s Bluff, S.E. of

TELINOLS seeeteh prt les 9095) elas
Other Material Examined: On
Cypress Swamp (Deer Pond), Johnson

islet ea isl Crane, 91-69, ILLS

Gorevill, Johnson County,
Crane 12-1-69, ILLS 42170.
decayed wood, Elvira
County Illinois, 22 May
42171.

Key to the Species of Codinaea Maire*

1. Conidiophores synnematous ......... C. obesispora (3)

Conti da ODNOneSHMOnenema cousin. sisrcisisiciclele sicleicis + + 01s sl eless sane
Zs Piva loconid Lamwit nese UULAC mets crclescholeie ts cherelelots otslekete erecennes
Pntaroconid ramwschOUGMSeCULACEI cr culstsisisiclietslc sloisiere siete sme’
B Phialoconidia with a setula at eachvend 2... ....e-+n0
Phialoconidlamwichiel api Ca larse Cu lasts cle seis lotta ice leleaee

4, Setula definite, 13-15 um long ..... C. unisetula (7)
Setula apiculate eeoseeeoeeeeweeeepeeeeweeeseseeeeeeeoeeevseeee see @ 5
5. Phialoconidia 14-19 x 3.3-4.4 um. C. illinoensis

Phialoconidia 20-30 x 4.0-5.5 um ... C. apiculata (6)
b PHPAMOCOM MOAT ARSE DLOUEC iW elels siclstcl sisicle.ctsisie.e sie eitte oiebetetaioieioue,

Rita bOCOnT di amnONn=SEDLALCmrleisse cles sic sisle cs cleiets slo oie sss uec
iemeesetulae <s3.0 um long 9--.<....+-.+ GC. brevisetulan( 3)

SOUILGCM aE SOmIMmLONE ahs ts otels ote leceretetstenars cleleiescre steeretereremn
8. Phialoconidia 3-septate .......... C. matsushimae

Phialoconidia l-—septate .ccccccossesccccovssvvecses
9. PAC ACMULOSEN Ciisic cise cisis sie boc areieie ieee eterelere oie cicieleie eietersteumnlU

Ser AGmaDSeNbUa. .elsisc.c es cle cisicle o\c eis cleie ss oleveleieiele «isles efemitii
10. Setae simple or branched, sterile or fertile;

Hale

CONTI aan d—al faeXeel | Ome o) ll Mace

Chaetosphaeria dengleyae anam. Codinaea (3)

Setae simple,

apex fertile;

conidia 13-20 x 2.5-3.5 um C. novae-quineensis (5)

Conidiophores

1-3-septate; conidiogenous cells

MSUalVeCenMitia laste cscs seis sa Se DL ac ama)

Conidiophores

up to 20 septate; conidiogenous

cells usually intercalary ....... C. eucalypti (11)

*Numbers in parentheses refer to literature cited.

426

12.
13.
14.

15.
16.

17.
ile,

19.
20.

AL fs

22.
o3e

24,

25.
26.

eT.

28.
29.
30.

Bae

Setae present
Setae absent .......
SCCAGaLERUCILES” 6 pisie & ciercls os 04.0) 0) she sieis clelcvois\e crelels
Setae sterile .....
Phialides from node-like hyphae

on setae
Phialides from base of setae
Setaeh<e250mims longi. 1. cs. sec epakKialensismaa)
Setacs>w250 Uma LON Rass alstcis com cic stele cciete rented er here mL O
Setae slightly geniculate at apex ... C. fertilis (3)
Setae straight, cyclindrical to gradually

Caperingsaceapex tei. ole e ec views Cemascam Camus)
Setulae of unequal length ........... C. aristata (4)
Setulae equal in length ..cccccccccccccees
Setutacv5-9 WmulOne@ae vse le eels ocr Dr Evanicam ain
Setulaepwup .UOuil « Onw Mal ON it ote ols cuss ecko lelelel oie cieteietetecetels

Chaetosphaeria pulchriseta anam. Codinaea (3)

Setulae’<n6 mil Ongawnc. c acts cele csisle trele cc cuterencrsirecmece
Setulaed 76 imrvoniers stele stele ste clove ate shelele c-o7e deceit tet temicnl
Phialoconidia ji=4 7 )umilong sc. <--> 1 ComDAnVaAmC 3)
Phialoconidia 17-23 um long ......... C. vulgaris (3)
Phialoconidia with acute apex,

eooeoevevreecee oe eere eee eee ee eee ee oe oe Lad,

gonytrichodes (10)

espe 0016, 6 cole elec eleraie cre .o MELO

eeoeeeoeeeeseftfeeeseeeee @ @ @ C.

ooo oe 8 @ @ 1

Phialoconidia

14-19 um long ....

Phialoconidia
Phialoconidia
Phialoconidia
Phialoconidia
Conidiophores

and ornamented ..ccccseeseeveee Co

Conidiophores
Phialoconidia

with bluntly pointed apex,
sessecsersoeceovce Cs Simplex: (35)
SCPCAL!S covcccccccvervsersescsescecees OF
NON=SCDtLALEL <.c.0 1c cies creretenele cit 6 clets ele seeeO
B=SEptate tes cece seme map LCdl Smo
T~Septateiic sc cists oles cc cistere seeleteiote aioe samc
branched

glauco-nigra (3)
SIMPLE And “SMOOUNete sietctetetele cic cl etetereiatel aac
10s 169 Xt 225=5. 5 msec eC ena Lam)

Phialoconidia 20-24 x 2.0-2.8 um ...... C. setosa (3)
Conldlopnores] Drancnedigs steterct ele cletsiclelc’c cleteleteretere! erence
Conidiophores simple @eeoeseeoeeeeseese@eeeeseerpeeeeeeeet eee 6 @ @ @ 28

Phialoconidia
Phialoconidia

20=26 um long ......... Co hughesi ind)
10—15 um long eoeeevoeveneeeeveee eee eee 8 &

Chaetosphaeria callimorpha anam. Codinaea (3)

Phialoconidia
Phialoconidia
Phialoconidia
Phialoconidia

SCGONE ya CUIVECS « cicicleisie c1e.sipis sieteisiete omc
straight or slighly curved ......-.. 30
Sigmoid ............ C. lunulospora

botuliform’......... C. botulispora’ (3)

Phialoconidia cylindrical ™..... C. ~ eylindrospora Gp)
Phialoconidia tapering towards base Mtn eens)

Phialoconidia

abruptly tapered

AL DAS] ccccecccccssccoesesceveees Co Clavulata (2)

Phialoconidia

gradually tapered at base .........6.

Chaetosphaeria talbotii anam. Codinaea (3)

427

Literature Cited

webs sao. 5. 1976. More dematiaceous Hyphomycetes.

Commonwealth Mycological Institute, Kew, 507 pp.

- Gams, W. and V. Holubova-Jechova. 1976. Chloridium

and some other dematiaceous Hyphomycetes growing on
decaying wood. Studies in Mycology No. 13, Baarn, 99
Pp.

Hughes, S. J. and W. B. Kendrick. 1968. New
Zealand fungi 12. Menispora, Codinaea, Menisporopsis.
New Zealand J. Bot. 6: 323-375.

Maire, R. 1937. Fungi Catalaunici, Series alterna.
Contribution a 1’etude de la Flore Mycologique de la
Catalogne. Publcions Inst. bot. Barcelona 3: 128
Pp.

Matsushima, T. 1971. Microfungi of the Solomon
Islands and Papua-New Guinea. Pub. by author, Kobe,
the) We

Matsushima, T 1975. Icones Microfungorum 9 a
Matsushima Lectorum. Pub. by author, Kobe, 209 pp.

Morgan-Jones, G. and E. B. Ingram. 1976. Notes on
Hyphomycetes XV. Two new species of Codinaea.
Mycotaxon 4: 504-509.

RIPOZyYNSKis Ka tA. sand S. D. Patil. 1970. Some
setose Hyphomycetes of leaf litter in south India.
Ganadiamtd .meeBOU. P4056 507—50 16

Reddy, S. M. cheyel | Sr S. 1978. A new species of
Codinaea Sydowia 30 (1-6): 186-188.

Shearer, | Ce A. Andee ot BOL ANC mero (clea Ul e1ero fy
the Chesapeake Bay and its’ tributaries. I. Patuxent
River. Mycologia 63: 237-260.

Sutton, Bb. .Cs and’. 6S. Hodges, 91975. Eucalyptus
microfungi: Codinaea and Zanclospora species from

Brazil. Nova Hedwigia 26: 517-525.

MYCOTAXON

Vol. XIII, No. 2, pp. 428-430 July-September 1981

A NEW SPECIES OF DACRYOPINAX FROM BRAZIL
B. Lowy

Botany Department, Louisiana State University
Baton Rouge, LA 70803

Dacryopinax maxidorii Lowy, sp. nov.

Fructificatio in humido aurantiolutea, elastico-
gelatinosa, pileata, 4.5 cm alta ad 5 cm lata; sicca
cornea, ferrugineo-brunnea; caulis elongatus, firme
radicatus, conspicuus, crassi-venatus; in sicco dense
albido-tomentosus; apicis alte ramosus, deinde poly-
cephalus, lobuli flabelliforme ad 450 um crassis cum mar-
gine crenulata; hymenio glabrum, unilaterali, inferiori,
pauci pilosa; probasidia cylindraceae, aseptata, 30.0-36.5
X 4.5 um, metabasidia furcata, aseptata, bisterigmata;
hyphae enodosae, 2.0-3.0 um diam; basidiosporae leviter |
curvulo-cylindraceae vel subovoideae, uniseptatae, in cu-
mulo aurantiaco-lutea, (7.0-) 8.0-10.0 (11.5) X 4.5-5.0
um, per promycelium aut conidia germinantes; conidia sub-
spherica, 1.5-2.5 um diam.

Fructification when fresh rubbery gelatinous, orange-
yellow, stipitate, pileate, up to 4.5 cm in height, 5 cm
broad (Fie. 1). drying horny, rusty brown; marisinge: roma
broad, elongated, firmly rooted, sterile stalk; thick
veined, densely whitish-pilose when dry, with pilosity
diminishing upward; deeply branched apically, frequently
becoming polycephalic with broad, veined, flabelliform
lobes + 450 um thick, often with crenate margins;
hymenium unilateral, inferior, producing a dense palisade
of basidia; abhymenium thinly covered with hyaline, cylin-
drical to slightly inflated, unbranched hairs, unicellular
to sparsely septate, arising from a layer of irregularly
inflated hyphae; probasidia cylindrical, unicellular,
30.0-36.5 X 4.0-4.5 wm; occasional cylindrical, slender,
unbranched dikaryophyses 20-25 X 2.5-3.0 um; metabasidia
furcate, bisterigmate; hyphae without clamp connections,

429

2.0-3.0 um diam; basidiospores slightly curved-cylin-
drical to subovoid with narrowed apices and prominent
apiculus, with a single, thick, central septum, (7.0-)
8.0-10.0 (-11.5) X 4.5-5.0 um, producing subspherical
conidia 1.5-2.5 um diam, or germinanting by germ tube.

Holotype, Brazil. Lowy 190 BR (TYPE), km 405 Manaus-
Porto Velho road, Amazonas, 16-I1X-1980.
Leg. 8B. Lowy, D. Coelho. On unidentified log.

This species is named for Maxine and Doris Lowy, enthu-
siastic collectors of tremellaceous fungi.

Figure 1. Basidiocarps of Dacryopinax maxidorii
with scale shown in millimeters.

In gross morphology the new species somewhat re-
sembles D. indacocheae Lowy (1959), except for the lat-
ter's stubby stalk, thinner lobes, and brownish to earth-
colored pigmentation. However, the long stalk, bright
carotinoid pigmentation, and flabelliform lobes of the
new species places it closer to D. martinii Lowy (1971).
The basidiospores of these three species are predominantly
l-septate, but D. maxidorii shares with D. indacocheae
somewhat curved-cylindrical spores with a single, thick
septum whereas the spores of D. martinii are character-—
istically thin septate.

430

The

following revised key includes the seven known

species of Dacryopinax occurring in tropical America.

Basidiocarp slightly expanded apically; pileus
simple r,orvbranched (en) (Pete Woe CaN mer eeee sy ut nee 2
2. Stipe tomentose; basidiospores becoming

septate, en. ea. oe I D. spathularia
2 obi pemno tu coment ose: Reet deepens: becoming

S=SeP Cates eer Uh ie ere eee: ; Deedennisit
Basidiocarp broadly expanded apically; pileus gross-
ly lobed to ftlabelln form onpepolycephalice.. =... 93
3b. slongestipmtateseplleuse:)abel 1. Orn: ee ee
3. Short stipitate to substipitate; pileus not

Flabelli form. ges ean ee ells Eh cre ode ee een aE
4, Abhymenium and stipe smooth; dried pileus thin,

fraci lev wen. ; Dinas Gd nda
4. Abhymenium and Eoime miteeer Farid pileus thick,

LODUS ET wi pee omen a ens eerie Detmas ident
5. Hymenium coarsely Penis yellow when

bresh yt mass ee: D.- yungensis
5. Hymenium sAereiel va ec when fresh . 6
6. Pileus spreading-orbiculate, undivided, drying

dark brown to blackish. . ; : D. elegans
6. Pileus foliose, SSP Toned drying light

Jopeeymaaie isos (ie Pa : D. indacocheae

ACKNOWLEDGEMENTS

Thanks are due The New York Botanical Garden, the
Instituto Nacional de Pesquisas da Amazonia (Manaus,
Amazonas, Brasil), and Louisiana State University (Baton

Rouge,

LA), under whose sponsorship the field work was

completed as part of the Projecto Flora Amazonica.

LOWY sab.

LITERATURE CITED

1959. New or Noteworthy Tremellales from

Bolivia. Mycologia 51: 840-850.

1971. Tremellales. Flora Neotropica. Mono-

graph 6, Hafner Publishing Company, Inc. New York.

MYCOTAXON

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MYCOTAXON

AN INTERNATIONAL JOURNAL DESIGNED TO EXPEDITE PUBLICATION
OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS

Volume XIII October-December 1981 NO

CONTENTS

Studies in the lichen genus Psoroma I: Psoroma tenue and
POPC CLIAINOME WI 6 6a ass oS ae od ele e's A. HENSSEN AND B. RENNER 433
An undescribed pleomorphic species of Codinaea... NORIHIRO TOYAZAKI 450
The psilopezioid fungi. VIII. Additions to the genus Pachyella.
DONALD H. PFISTER AND FRANCOISE CANDOUSSAU 457
Pithomyces pavgitt, a new combination for Trtchocladtum pavgtt

and Pithomyeces funtculosa...... MARY E. PALM AND ELWIN L. STEWART 465
Validation of the Herpomycetineae and Herpomycetaceae in the

SPUD ERE ALCS 2505s pacn, PURSE Taree ete, wee it UG ake Si ISABELLE I. TAVARES 469
Erynta (Zygomycetes: Entomophthorales): validations and new

PRES Wh et A ESP ran Bit Sie Aisa sats umes Ie Whee at alti nie foegia.y ae wie RICHARD A. HUMBER 471
A review of the nonentomogenous Entomophthorales... BRUCE E. TUCKER 481
Erynta (Zygomycetes: Entomophthorales): emendation, synonymy,

Mane Transfers. 3 ih sila. aes RICHARD A. HUMBER AND ISRAEL BEN-ZE'EV 506
eS LINTOS Aro Wo iia eee ia eee ahs ee G. L. HENNEBERT 517
Notices

Pere DI ce oCTence, CONPTESS., Klis is oe) oh Cite onin date whine se ea wiea An ples 525

IMC: A SA ECOS DS TBA ANAT o suuia vy novi Gerns fe elas bow Bhs pi wel ee! ay date Sova aby 525
PEE a ENG pte seed i oa og a bs oe ae Wee cp aie, She Wg AhG Gra sino a ine Bia © et bMeeW tale 526
MEE TROIS GL LCNCMOLAXAGL iui. Se ice Oo mk lh oye Mieipid ote Wide Wego: 8 528
ewen.ON publication dates 712 (2), 13(1) ;.and 13. (2). oo. elk. 534
See ee AN ee Te, SMU Ae lia tis Who Re es iehteed oe Wo ee Re ele meen 535
RE DR eae ias Becht EG tole th hy LCL ely) TUE: plate Vepa aia aA ba Sep Epo Ro edodel ic ovpnohnnc of JD

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MY COTAXON

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Pete ALTE NO. 3, pp. 433-449 October-December 1981

STUDIES IN THE LICHEN GENUS PSOROMA I
PSOROMA TENUE AND PSOROMA CINNAMOMEUM

A. HENSSEN and B. RENNER

macnberevehaBiologie ‘der Universitat Marbung
D-3550 Marburg/Lahn, Germany

SUMMARY

The new species -Psoroma tenue Henssen belongs to
the P. hypnorum-group and is most closely related
to P. cinnamomeum. Two varieties are described. P.
tenue var. tenue is restricted to the Southern He-
misphere; the lichen has been collected in Tierra
del Fuego and the South Shetland Islands. P. tenue
var. borealis Henssenis a wide-spread northern ta-
xon; localities are given for the Alps, Scandina-
via, Iceland and North America. P. tenue is a spe-
cies of snowbanks, glacial till and tundral heath.
The lichen contains porphyrilic acid methyl ester
and pannaric acid. The occurrence of these meta-
bolites in other members of the P. hypnorum-group
is mentioned. A short description of anatomical
structures in P. cinnamomeum is included.

INTRODUCTION

The lichen genus Psoroma Michx, a member of
the Pannariaceae, occurs predominantly in the
Southern Hemisphere, in South America and New Zea
land. The type species P. hypiorum (Vahl) S.F.
Gray (Synonyms: P. bryanttt Dodge as "Bryantt" in
imac aeAmMersrM1 crOoscop. Soc. 842511719055. holos
type: hb. Dodge; and P.follmannit Dodge in Trans.
aie wee Croscopm cOC. 164: 51.0,)a196500 1SOLY pe has
SEL) only is known to be world wide distributed.

434

Our studies in Psoroma are mainly restricted
to South American species, based on collections
made by A. Henssen and G. Vobis in southern
Chile and Argentina in 1973/74. P.W. James who
was planning a monographic treatment of the
genus encouraged us to collect Psoroma species
intensively. The series of Psoroma papers, pub-
lished in part together with P.W. James and K.
Marton, may be regarded as preliminary studies
LOX a world monograph ings thtseini Lid apapon we...
Henssen is responsible for the taxonomic treat-
ment; and B. Renner, for the identification of
the lichen substances.

InvLeS CuUrrentecirCcumMSeHl pELOnge se rCngm smd
heterogenous genus, and affinities of certain
species to Pannarta Del. in Bory have been sug-
gested (James & Henssen 1976, J¢rgensen 1978).
The species of the P. hypnorum-group, for example,
form a distinct assemblage of terricolous lichens
of cold temperature or subalpine / alpine areas.
They may be more closely related to Pannarta
pestzotdes (G.H. Web.) Trevis. than to the corti-
colous / saxicolous, squamulose or lobate species
of Psoroma with which they are at present allied
(James & Henssen 1976).

Thesnew) Species WP cucu De lLOnGgSme tL Omstlicomne.
hypnorum-group. The outstanding character of the
species is a pattern of lichen substances not
previously known) to soccuGn in Vesorengus.. me ne
specimens of P. tenue so far studied always con-
tain pannaric acid and the new compound porphy-
rilic acid methyl ester and some specimens con-
tain an unknown substance (U,) (Renner et al.
1981). Specimens from the Northern Hemisphere
differ slightly in morphology fromethe “southern
hemispheric ones and are included in the variety
borealts.

In P. hypnorum, no lichen substances were
found by TLC (J¢grgensen 1978). We confirmed this
observation in the type specimen (C) and in the
types of the synonymous species P. bryantit (hb.
Dodge) and P. follmanntt (KASSEL) as well as ina
number of our specimens from northern and southern

435

localities. All of these specimens were similar
with regards to external and internal morphology.
However, certain specimens, deviating to some
extent in morphology, were found to contain one,
two or all three of the compounds described from
P. tenue. These specimens need further taxonomic
Pavestigation. Porphyrilic acid methyl ester and
mennaric acid also occur in the new P, rubro-
marginatunP. James & Murray (Henssen et al. in
manuscript) , and the unknown (U, ) occurs ina
South American specimen of P. paleaceum (Fr.)
Nyl. (2? P. hirsutulum Nyl.) (Henssen 24353g: MB).

Within the P. hypnorum-group, several patterns
in thallus anatomy may be recognized. P. tenue
corresponds to P. ctnnamomeum in external and
internal morphology of the thallus and cephalodia:
The two species, therefore, are regarded to be
closely related. Although the South American
species P. etnnamomeum was well - described by
Malme (1952), additional information obtained by
modern methods is included in the presented paper.

MATERIAL AND METHODS

MATERIAL. The abbreviations of herbaria follows those
given in Index Herbariorum; hb. Dodge means the private
herbarium of C. Dodge, Burlington, Vermont, U.S.A., and
hb. Henssen means the private herbarium of the senior
author.

Psoroma cinnamomeum: Argentina, Tierra del Fuego,
Rio Grande, 1896, Dusén 115 (holotype: S); Cabo San Pablo,
on plant debris in a tundra heath, overgrowing a dead
liverwort, 1973, Henssen & Vobis 24448t (MB). P. hypnorum
(selected specimens): Norway, Vahl (lectotype by Jgérgensen:
(C). Sogn og Fjordane, Nordfjordeid, on rock overgrowing
mosses, 1959, Henssen 5339 (hb. Henssen). - Finland, Lap-
-ponia enontekiensis, Toskaljarvi, 1955, Henssen 6145 (hb.
Henssen). - Argentina, Tierra del Fuego, Glaciar Martial,
on rock overgrowing mosses, 400m s.m., 1973, Henssen &
Vobis 24412a (MB); Laguna Escondida, Henssen & Vobis

24439h and 24460d (MB). - Antarctic Peninsula, Refuge Is-
land, 1940, Bryant 30-2 (holotype of P. bryantii: hb.
Dodge). - South Shetland Islands, Greenwich Island, Bahia

Soberania, on mosses, 1963, Follmann 11843 (isotype:

436

Fig. 1. Psoroma tenue var. tenue (paratype), young and

mature densely aggregated apothecia; scale = 1 mm; drawn
by H. Becker.

of P. follmannii: KASSEL). P. paleaceum: Argentina, Tierra
del Fuego, between Ushuaia and Lapataia, on rock overgrow-

ing mosses, 1973, Henssen & Vobis 24353g (MB). P. tenue:
see below.

437

MORPHOLOGY. Samples were sectioned by freezing micro-
tome and the sections mounted in lactophenol cotton - blue.
Measurements of spores and anatomical structures were made
from permanent preparations; measurements of gross morpho-
logy on air-dried specimens. For studying the ascus
structure, KI, solution was added directly to the squash
preparation.

CHEMISTRY. Crude lichen extracts were chromatograph-
ed in solvent systems A, B, C of Culberson (1972), and D,
E of Renner (1980). The methods of mass spectrometry and
UV/VIS spectroscopy were used for the identification of the
lichen substances; details are given in Renner et al.
WEOGI)=

TAXONOMIC PART
Psoroma tenue Henssen, sp. nov. var. tenue

Hab) eb Ole cA Ore a | MismalalOny yen lOO ATe . ast Lue—
Luresor the ascocarp, Hige4A-D.

DIAGNOSIS. Thallus granulatus vel subsquamulosus,
pallidus vel fuscescens, e verrucis formatus semiglobosis,
circiter 0.2 - 0.4 mm latis vel e squamulis plus minusve
adscentibus, .0-45— 078mm) latis, crassis,) dispersis vel
subcontiguis. Thallus corticatus, hyphae medullosae reti-
culum formantes, alga symbiotica laete viridis, ad fami-
liam Chlorococcaceae pertinens. Cephalodia globularia vel
ramosa, plus minusve aggregata, atropurpurea, corticata,
alga ad Nostoc pertinens.

Apothecia usque ad 2 (-3)mm lata, disco plano vel sub-
plano, atrofusco, margo thallinus vulgo pallidus, tenuis,
subtiliter crenulatus. Hymenium 95-120 (-150)um altum,
Asci subcylindrici, 80-110 x 8-12ym, 8-spori, annulo amy-
loidea in apice asci incrassato, Sporae eseptatae, inco-
lores, polymorphae, ovoideae vel subfusiformes, 19-24 x
(5-)7-10 (-12)um, superfice verruculosa. Pycnidia minuta,
circiter 0.1-0.16 mm lata, conidiophora brevicellularia,
conidia terminalia et lateralia formantia. Conidia bacil-
bi formia, Circiter 2-4 x 1 um.

Chemistry: thallus and apothecia contain porphyrilic

acid methyl ester, pannaric acid, in the holotype the un-
known substance (Uj) was found.

438

439

Holotype: Argentina, Tierra del Fuego, Parque Nacional
de la Tierra del Fuego, Laguna Escondida, on loamy soil
near the lake shore overgrowing Cephaloziella and plant
debris, at c. 270m s. m. 1973 Henssen & Vobis 24439b (MB);
isotypes: BM, LPS. Paratype: near the type locality at
Hosteria Petrel, on soil overgrowing plant debris ina
stand of Nothofagus at the lake shore, 1973 Henssen &
Vobis 24460w (MB).

FURTHER SPECIMEN EXAMINED: South Shetland Islands,
Greenwich Island, on earth and plant debris, 1963, Follmann
14065 (KASSEL) .

Thallus of pale or more rarely brown colour, dull,
composed of dispersed or aggregated minute granules and
squamules, fastened by rhizoidal hyphae with colourless
or brown thick walls. Granules up to 0.4 mm, squamules 0.4
- 0.8 mm broad, more or less ascending. Thallus sections
(2559>)) 150-250 (450) um high, at least on the upper surface
distinctly corticated. Upper cortex in young thalli compo-
sed by thin-walled cells (Fig. 3A), later on the walls
gelatinizing, and the lumina in the gelatinous matrix up
to 18um broad, connected by pits (Fig. 3B). Lower cortex
composed of isodiametric or elongated cells (Fig. 3C).
Medullary hyphae (2.5-)4-6um thick, orientated reticula-
tely and enclosing single or several algal cells. Phyco-
biont a member of the Chlorococcaceae, cells c.6-10pum
broad, distributed throughout the thallus but more numer-
ous in the upper part. Crystals of lichen substances de-
posited in clusters in the medulla.

Pig. 2. Habit photographs of Psoroma species. A-B, P. tenue
var. tenue (paratype); A, lichen overgrowing plant debris;
B, enlarged apothecia. C, P. tenue var. tenue (holotype) ;
dark colored, aggregated cephalodia between thallus
squamules and apothecia. D, P. cinnamomeum (holotype). E-F,
P. tenue var. borealis (holotype); E, mature apothecia; F,
cephalodia (arrowed) between squamules. G, P. cinnamomeum
(holotype) , crowded apothecia surrounded by a thick thal-
line margin; the black apothecia belong to a Badicia
species. H, P. tenue var. borealis (hb. Henssen 6130),
Squamules and apothecia. A-H, scale = 1 mm.

440

441

Cephalodia granulary or coralloidly branched, dark
purplish brown, singly or forming clusters up to 0.8 mm
(Fig. 2C). Cephalodia corticated, cortex 4.5-12yum thick,
formed by 1 to 2 rows of cells; medullary hyphae orientated
in a network in young stages, later forming pseudoparen-
chyma (Fig. 5E). Alga a Nostoc species.

Apothecia dispersed or closely aggregated, up to 2(-3)
mm broad, disc flat, dark brown, surrounded by a pale,
finely crenulated margin (Fig. 1, 2A, B). Hymenium 95-120
(-150)um high; hypothecium - layer including the ascoge-
nous hyphae - 20-25(-50)um; subhymenial layer of varying
thickness, frequently extending into a stipe of adglutinat-
ed hyphae with enlarged cells towards the base. Margo thal-
linus corticated, basal part 100-120um broad, composed of
radiating hyphae with broad, up to 25um wide cells; joint
walls between the cell lumina 3.5-6um thick (Fig. 4A, B).
Crystals of lichen substances in clusters in the medulla
of the margo thallinus.

Pycnidia inconspicuous, in sections 0.1-0.16 mm large.
Conidiophores branched and anastomosing, conidiogenous
cells short, the conidia terminally and laterally produced.
Conidia rod-shaped, c.2-4 x lpm.

Psoroma tenue var. borealts Henssen var. nov.

Poot eee, (so challus anatomy, /f19. 3D-F; seruc—
Lure sOL apothecium, Fig. 5C.

DIAGNOSIS. Differt a var. tenuethallo majori semper
distincte corticato et colore cinnamomeo vel luteofusco.

Chemistry: thallus and apothecia contain porphyrilic
acid methyl ester and pannaric acid, in the holotype and
in the specimens from Austria, Ost-Tirol and Canada the
unknown substance (U,) was found.

Fig. 3. Thallus anatomy of Psoroma species (microtome
Sections). A-C, P. tenue var. tenue (holotype); A, 1.s. of
young thallus, crystals of lichen substances indicated by
arrow; B, t.s. of upper cortex and algal zone; C, t.s. of
thallus adjacent to an apothecium. D-E, P. tenue var. bore-
alis (Henssen 22679g); D, 1.s. of young thallus; E, 1.s. of
a cephalodium at the base of an apothecium. F, P. tenue
Var. borealis (Henssen 23157x), t.s. of corticated thallus.
A-F, scale = 20 um.

442

443

Holotype: U.S.A., Colorado, Boulder Co., Niwot Ridge,
on snowbank, between 3300 and 3750 m, 1961, Henssen 13030b
(hb. Henssen). Paratype: Austria, Tirol. Samnaungruppe,
Zeblasjoch, on snowbank at 2450m, 1975, Henssen 22679g (MB).

FURTHER SPECIMENS EXAMINED: Austria, Ost-Tirol,
Matrei, Johannishitte, in glacial till at c. 2200 - 2400 mn,
overgrowing mosses, 1938 F. & Koppe (hb. Henssen 6130). -
Norway, S6r-Troéndelag, Dovrefjell, Knuthd, on snowbank at
1500 m, 1959, Henssen 4974 (hb. Henssen). - Sweden. Torne
Lappmark, Abisko. Nuolja, on snowbank at 1160 m, 1954,
Henssen 6127 (hb. Henssen). - Iceland, Bard, Kollsvik,
overgrowing mosses in a lichen heath, 1972, Henssen &
Kristinsson 23157x (MB). - Canada, Alberta, Banff National
Park, Peyto Lake, Dryas heath at 2150-2250 m, 1962, Henssen
& Cain 14471y (CAN, MB).

Thallus squamules up to 1 mm broad, brown or reddish
brown, thick; thallus distinctly corticated on the upper
and lower surface by isodiametric cells (Fig. 3D, F).
Apothecial disc brown and margo thallinus of the same
colour, or disc dark brown and margo thallinus slightly
paler. Crystals of lichen substances deposited within the
medulla of the thallus and apothecial margin or not.

Pie emecenuesvar. Lenue the lobes are smaller
Brae mevatemlOneca, is (Fig. 92) ,andvthe cortex is
usually less well developed since it is restric-
ted to the upper surface. Broad cortical struc-
tures on both thallus surfaces have been observed
in thalli closely attached to margins of apothe-
Slam fg. 56)r in such lobes the thallus corres=
ponds exactly to that in P. etnnamomeum (cf.
meGs. 3B with’ 5A).

The cephalodia occur either between the lobes
Or are producted in groups below the apothecia
(Puge038).) the alga in the cephalodia is a

Fig. 4. Apothecial structure in P. tenue var. tenue (micro-
tome sections of the holotype). A, median 1l.s. of young
apothecium, note the numerous algal cells in the stipe; B,
cortex structure in apothecial margin; C, upper part of
apothecial margin, crystals of lichen substances indicated
by arrows; D, ascus and spores. A-D, scale = 20 um.

species of Wostoc as is found in most Psoroma
species.

The margin of the apothecia develops in the
Same Way as in ‘Pannaria rubtginosa (Ach.) Bury
(Henssen 1969, 1981). The cortex in the margo
thallinus is differentiated in part by the thal-
lus cortex and in part by the medullary hyphae
growing up between the hymenium and the thallus
cortex (Fig. 4A, B). The cells formed by the me-
dullary hyphae are considerable smaller in size
(Fig. 4C, 5C). As seen from above the finely cre-
nulated margin surrounding the relatively flat
disc resembles that of certain Pannarta species.
In very old apothecia the margin may bear granu-
les and the disc becomes folded or medianly de-
pressed. Young developing apothecia are at first
urceolate.

The ascus wall and hymenial gelatine stain
blue in iodine, and an amyloid ring structure is
seen in the ascus apex of the same shape as that
Pound in 2. 2ypnorum (Keuck 1977).

hie emer vee var. tenue yellowish to brownish
crystals of lichen substances are usually depo-
sited in large amounts in the medulla of the
eoellus and apothecial margin. In var. borealis
Similar crystals have only been observed in the
specimen collected in Canada (Henssen 14471y ).
miese crystals are difficult to demonstrate in
the sections by black and white photography but
are rather impressive in microscopic examination
of stained permanent preparations. The varying
occurence of the unknown substance (U,) in both
varieties of the species is interesting.

Fig. 5. Anatomy in Psoroma species (microtome sections).
A-B, P. cinnamomeum (holotype); A, 1.s. of thallus; B,

_ cortex structure in apothecial margin; C, P. tenue var.
borealis (Henssen 22670g), apothecial margin; D, P. cinna-
momeum (Henssen 24448t), cortex structure in apothecial
Margin; &, ~. tenue var. tenue (holotype), 1.s. of cepha-
lodium; F, P. cinnamomeum (Henssen 24448t), 1.s. of cepha-
lodium. A-F, scale = 20 um.

446

Fig. 6. Psoroma cinnamomeum (Henssen 24448t), young apo-
thecia; scale = 1 mm; drawn by H. Becker.

PSOrOMma Lene mS GnanractenuZeds Dy eLnempEesence
of porphyrilic adic methyl ester and pannaric
acid’ in) correlation with, small, flatvanotnecia
surrounded by a thin evenly crenulated margin, by
dark purplish brown cephalodia, aggregated to
clusters, and by an upper thallus cortex formed
by periclinal hyphae with isodiametric cells.

In .comparisoni toe. venue.) P. vetnnamoneumsnas
a Similar anatomy and the same type of cephalo-
dia but) differs eapartetrom stiews acwromeltecnen
substances, by having a very thick apothecial
margin in the young, strongly urceolate apothecia

447

and by the smaller lumina in the cells forming
Pies COLGCxK@ ine thesmargo thallinus (cf. Figs, 4¢
Pom Gawiltlimb, mi )ree ect nnanomeumn OCCULS in the
PameeDegioOns as) Ps venue Var. tenue. By the pale
So OUImMOms ies thialius tand athe abundant crystals
meapothecialior thallus sections, £. tenue var.
tenue is easily distinguished from P. etnnamo-
meum. Reddish brown specimens of P. tenue var.
borealts resemble P. etnnamomeum but the variety
is only known from the Northern Hemisphere at
this time.

ECOLOGY. P. tenue var. tenue has been collec-
ted on soil, long covered by snow, together with
Pe Aypnorum and P. pateaceum. P. tenue var. bore-
alts has been found, hitherto, on snowbanks in
glacial till and in tundral heath, where the li-
chen was growing together with Lectophysma ftn-
Po ieee merbivett 96 235 ox) (OLei meri urascens, (Ny!.)
Gyeln. (14471y) respectively, among other lichens.

Psoroma ctnnamomeum Malme 1925 P. 11, 12

The specimen studied in comparison to the ho-
lotype (Henssen 24448t) was growing on the same
type of substrate. The prominent apothecial mar-
gin is especially distinct in younger apothecia
(Fig. 1G, 6). The cortex was well developed in
all'sections studied. In the upper cortex the
cell lumina are embedded in the gelatinous matrix
of the cell walls. The lumina are roundish and
@onnected by pits (Fig. 5A). In young thalli the
cortex is formed only by periclinally arranged
hyphae, later on cells are added by vertically
growing medullary hyphae. In the lower thallus
cortex the cell walls are less strongly gela-
man Zed. tThe, cortex in the margo thallinus is
formed by radial or somewhat reticulately arran-
ged hyphae with thick walls and relatively small
Poin amiie LC .aeoB, )D)

The cephalodia are dark purplish brown and
form clusters between the squamules or they are
produced in groups below the apothecia. They are
surrounded by a brown pigmented cortex, composed
PeeOnewOL,sinupabt, two)rows Of cells (Fig. 5F).

448

In young stages the hyphae are thin and form a
network and later the enlarged cells become ag-
gregated into a pseudoparenchyma.

ACKNOWLEDGEMENTS

These studies were supported by a grant of the Deutsche
Forschungsgemeinschaft, which also provided a travel grant
for the field studies in South America. In Tierra del Fuego
we were guests at the Marine Biological Station in Ushuaia,
and the excursions were guided by Dr. S. Guarrera from the
University of La Plata. We are greatly indebted to him and
to Dr. Kthnemann for the profitable stay we had in this
remote part of the world. We wish to thank Dr, C. Dodge,
Burlington and the curators of the herbaria for the loan
of the type material. Mrs. G. Traute is thanked for skilled
technical assistance and Dr. Nash for reading and correct-
ing the manuscript.

ZUSAMMENFASSUNG

Die neue Art Psoroma tenue Henssen, eine Flechte auf
Schneebdéden und in Tundra-dahnlicher Vegetation gehort zur
Psoroma hypnorum-Gruppe und ist am ndchsten mit P. cinna-
momeum Malme verwandt. Morphologie und Anatomie der beiden
Arten werden verglichen. Sie stimmen in der Wuchsform, der
Struktur des Lagers sowie in der Gestalt und Anatomie der
Cephalodien weitgehend tiberein. Zwei Varietdten der neuen
Art P. tenue werden unterschieden. P. tenue var. tenue ist
auf die stdliche Hemisphdre beschrdankt und wurde bisher in
Tierra del Fuego und auf den South Shetland Islands ge-
sammelt. P. tenue var. borealis ist anscheinend eine weit
verbreitete Flechte der nérdlichen Hemisphare; Fundorte
werden ftir die Alpen, Skandinavien, Island und Nordamerika
angegeben. Beide Varietdten von P. tenue enthalten den
neuen Flechtenstoff£f Porphyrilsaduremethylester und Pannarin-
sadure, Sowie in einigen der untersuchten Proben zusdtzlich
eine unbekannte Substanz (U,). Das Vorkommen dieser Stoffe
in anderen Arten der P. hypnorum-Gruppe wird erwahnt.

Psoroma bryantii Dodge und P. follmannii Dodge werden
als Synonyme Zu P. hypnorum (Vahl) S. F. Gray gestellt.

449

RE RE RENGES

Culberson, C. F. 1972. Improved conditions and new data for
the identification of lichen products by a standardized
thinglayerschromatographic method. J. «Chromatogr. 72:
ie 7

Henssen, A. 1969. Die Entstehung des Thallusrandes bei den
Pannariaceen (Lichenes) mit einer generellen Diskussion
uber die Entwicklung lecanoriner und biatoriner Flech-
tenapothecien. Ber. Dtsch. Bot. Ges. 82: 235-248,

Henssen, A. et al. 1981. The Lecanoralean Centrum. In
Ascomycete Systematics The Luttrellian Concept (D. R.
Reynolds ed.): 138-234. New York, Heidelberg and Ber-
lin: Springer-Verlag.

James, P. W. and HensSen, A. 1976. The morphological and
taxonomic significance of cephalodia. In Lichenology:
Progress and Problems (D. H. Brown, D. L. Hawksworth
and R. H. Bailey, eds): 27-77. London, New York and
San Francisco: Academic Press.

Jgrgensen, P. M. 1978. The lichen family Pannaricaceae in
Europe. Opera Botanica 45: 1-124. NFR: Stockholm.

Keuck, G. 1977. Ontogenetisch-systematische Studie tber
Eridoderma im Vergleich mit anderen cyanophilen Flech-
tengattungen. J. Cramer, Bibliotheca Lichenologica
WO Lem own Vaduz.

Malme, G.O. 1925. Die Pannariazeen des Regnellschen Herbars.
AG DOC a Z0AENOgs 2 .l—2 3.

Renner, B. 1980. Untersuchungen zum Einflu8 der symbio-
tischen Alge auf den Stoffwechsel und die Struktur des
Flechtenlagers. Dissertation, Philipps-Universitat,
Marburg (microfiche).

Renner, B., Henssen, A. and Gerstner, E. 1981. Pannarsaure
und Porphyrilsduremethylester - Sekundarstoffe der
Flechtengattung Psoroma. Z. Naturforschung (in press).

MYCOTAXON

Vol. XIII, Nox. 3, pps 450-456 October-December 1981

AN UNDESCRIBED PLEOMORPHIC SPECIES OF CODINAEA
Norihiro TOYAZAKI

Public Health Research Institute of Kobe City, Minato-
jima-naka-machi 4-chome, Chuo-ku, Kobe 650, Japan

AND
Shun-ichi UDAGAWA

National Institute of Hygienic Sciences, Kamiyoga
l-chome, Setagaya-ku, Tokyo 158, Japan

In the course of an investigation of airborne fungi in
Kobe, Japan, a hyphomycete was obtained in pure culture
which proved to represent a previously undescribed species
of Codinaea Maire. When grown on a variety of media such
as cornmeal agar, malt extract agar, oatmeal agar, potato-
carrot agar and potato-dextrose agar, sporulation is pleo-
morphic: lt produces (a) polyphialidicysdarker wa thickoas
walled, and regularly septate conidiophores and allantoid,
hyaline, 3-septate, and terminally setulate conidia charac-
teristic of Codinaea, and (b) monophialidic, paler, and
less septate conidiophores with fusiform non-septate con-
idia somewhat suggestive of Chloridium. Due to the latter
features, the isolate superficially resembles Chloridium
codinaeoides Pirozynski (1972), which produces monophiali-
dic conidiogenous cells and narrowly fusiform-obclavate,
aseptate conidia.

A recent account of the morphology and taxonomy of the
genus Codinaea was given by Hughes and Kendrick (1968), who
recognized 13 species and four anamorphic forms of Chaeto-
sphaeria (Sphaeriaceae) and described in detail numerous
collections from New Zealand. Subsequently the following
13 species have been added to the genus: C. maharashtrensis
Pirozynski § Patil (1970), C. novae-guineensis Matsushima
(1971), C. eucalypti Sutton § Hodges (1975), C. septata
Sutton §& Hodges (1975), C. apiculata Matsushima (1975), Cc.
Tunata Matsushima (1975), =CosbritannicasM «= Byeoll ts eUuLoyOl.
C. hughesii M. B. Ellis (1976), C. cylindrospora Morgan-
Jones & Ingram (1976), C. unisetula Morgan-Jones §& Ingram

451

(1976), C. clavulata Holubov4-Jechova (1976), C. parkhalen-
sis S. M. & S. S. Reddy (1977), and Codinaea anamorph of
Striatosphaeria codinaeophora Samuels §& E. MUller (1978) (as
'"S. codinaeaphora'). Dictyochaeta fuegiana Speg., the type
species of the genus Dictyochaeta Spegazzini that was con-
sidered as a possible earlier name for Codinaea (Hughes and
Kendrick, 1968), has been re-described by Godeas et al.
(1977). However, there are too few diagnostic features
available on the type to properly define the type species
of that genus.

Since none of the fungi of this genus are pleomorphic,
the fungus from Kobe has been described here as a new
species.

Codinaea dimorpha Toyazaki & Udagawa, sp.nov. (Figs. 1-6)

Solonvacmineagaromucormmealy slentcecrescentes wm itciuts—
simae, ex mycelio vegetativo submerso compositae; hyphae
aeriae sparsae; conidia abundantia, aggregata, brunneo-
grisea; reversum incoloratum. Coloniae in aliis agaris
communibus celerius crescentes, planae vel rugosae, tenues,
penitus auratae vel aurantiacae.

Mycelium immersum vel semi-immersum, ex hyphis hya-
linis vel subhyalinis, ramosis, septatis, 1-3 um dian,
laevibus compositum.

Boudacmsteri cSeansentes:

Structurae conidiogenae macronematae, dimorphae: (a)
polyphialidicae et (b) monophialidicae. (a) Conidiophora
ex hyphis repentibus superficialibus orientia, singula vel
2-3 aggregata, simplicia, brunnea, cylindrica, 200-280 x 6-
8 um, laevia, incrassata, omnino diametro uniformia, apicem
versus pallescentia, recta vel flexa vel plerumque genicu-
lata, usque 6-8 septata, superne successive elongascentia.
Cellulae conidiogenae elongatae, cylindricae, vulgo usque
ad 40-60 um longae, polyphialidicae, raro discretae, deter-
minatae, pallide brunneae vel subhyalinae, multis collari-
bus conspicuis praeditae; collaria infundibuliformia, 3-4 x
1.5-2.5 um, subhyalina vel pallide brunnea, postremo
effracta. Conidia semi-endogena, acropleurogena, hyalina,
allantoidea, ellipsoidea vel oblonga, parum curva, 22-28 x
7-8 um, 3-septata, laevia, ad basim interdum hilo incon-
spicue et utrinque setula singula, simplici, hyalina, 5-6
um longa praedita, in massa mucida aggregata. (b) Conidio-
phora ex hyphis superficialibus singula oriunda, simplicia
vel interdum irregulariter ramosa, 2-3 phialides ferentia,
eyiindrica; 45-100 x 2.5-4 ym, hyalina vel obscure oliva-
ceo-brunnea, prope basin septata, laevia. Cellulae coni-
diogenae monophialidicae, integratae, determinatae, cylind-

452

ricae, superne gradatim angustatae, collari distincto 2-2.5
x 1-2 wm praedita. Conidia semi-endogena, acrogena, hya-
lina, cylindrica vel fusiformia, 10-18 x 1-2 um, parum
curva, aseptata vel raro uniseptata, utrinque plus minusve
truncata, in massa mucida aggregata.

Chlamydosporae terminales, obscure olivaceo-brunneae,
ellipsoideae vel pyxritormes, slacves,inCrassal ae, Ul on
6-10 um, aseptatae vel raro uniseptatae.

Holotypus: cultura NHL 2891 ex aere urbis Kobe in
Japonia,in I2.vi: 1980 (maeNw Loyazaki, 1sOlata |e inecol tece
tione fungorum "National Institute of Hygienic Sciences
(NHL), Tokyo, Japan."

Etymology: lat. dimorphus = having two forms, referr-
ing to the dimorphic conidiogenesis.

Colonies on cornmeal agar (CMA) growing slowly,
reaching 2.8 cm in diam after two weeks at 23 C, very thin,
vegetative mycelium largely submerged; white aerial hyphae
sparsely developed on the agar surface; conidia-bearing
structures abundantly produced in central colony areas,
"brownish grey'' (Kornerup and Wanscher, 1978; pl. 6-E-2);
reverse uncolored.) Coloniesson malt extract _ycascmcxtrace
agar (MYA) growing rather restrictedly, consisting of a
somewhat thick basal felt, wrinkled conspicuously, velvety
to floccose, producing abundant aerial hyphae, "pastel yel-
low'' (Kornerup and Wanscher, 1978; pl. 2-A-4); conidia
slowly developed, not affecting the colony appearance with-
im three weeks: sreyerse "Orange ma hormerupeancendisciic ia.
1978; pl. 6-B-7). Colonies on oatmeal agar (OA) or potato-
dextrose agar (PDA) growing more rapidly, reaching 3.0-3.6
Cm) in diam arter stwo-weekssaty25 UC, plane chil, .« cecum) ec. —
low" or "deep orange’! (Kornerup and Wanscher, 1978; pls. 4-
A-8, 5-A-8).

Mycelium immersed or semi-immersed, composed of hya-
line to subhyaline, branched, septate, 1-3 um wide, smooth-
walled hyphae which often swell to knots of pale brown to
brown cells up to 5.5-10 um wide.

Sterilessetgemiacking.
Conidia-bearing structures macronematous, of two
types; (a) polyphialidic, and (b) monophialidic. Type (a)
commonly developed on CMA and at later stages also on MYA,
OA and PDA. Conidiophores arising singly from superficial-

ly repent hyphae or in groups of two or three from hyphal
knots, simple, brown, cylindrical, 200-280 x 6-8 um, smooth
and thick-walled, uniform in width throughout, paler to-
wards the apex, straight or flexuous, often markedly genic-
ulate in the upper region, up to 6-8-septate, successively
proliferating with age at short distances. Conidiogenous

453

Fig. 1. Codinaea dimorpha (Ni Dez Soe
A. Conidiophores and polyphialides. B. Conidia(3-septate) .
C. Monophialides and conidia. D. Chlamydospores. (All

measurements are in um.)

454

Figs. 2-6. Codinaea dimorpha (NHL 2891).
2. Conidiophores. 3. Three-septate conidium. 4. Mono-
phialidic conidiogenous cell and 0-l-septate conidia. 5 and

6. Polyphialidic conidiogenous cells. (Bars in 3-6 = 10 um
and in 2 = 100 um.)

cells elongated, cylindrical, usually up to 40-60 um in
length, -polyphialidic, rarely discrete, determinate, pale
brown to subhyaline, each phialide in turn becoming exter-
nally scarred with the persistent remains of the collaret-
tes; collarettes funnel-shaped, 3-4 x 1.5-2.5 um, subhya-
line to pale brown, aged ones often broken off. Conidia
semi-endogenous, acropleurogenous, hyaline, allantoid, el-
lipsoid or oblong, slightly curved, 22-28 x 7-8 um, 3-sep-

455

tate, smooth-walled, sometimes with a basal hilum, provided
at each end with a single, simple, hyaline setula 5-6 um in
length, aggregated in a slimy mass. Type (b) developed at
an early stage on MYA, OA and PDA. Conidiophores arising
singly from superficial mycelium, simple or sometimes ir-
regularly branching into two or three stalked phialides,
cylindrical, 45-100 x 2.5-4 um, hyaline to dark olive-
brown, septate near the base, smooth-walled. Conidiogenous
cells monophialidic, integrated, determinate, cylindrical,
tapering gradually into a well-defined, terminal collarette
measuring 2-2.5 x 1-2 um. Conidia semi-endogenous, acro-
genous, hyaline, cylindrical to fusiform, 10-18 x 1-2 um,
slightly curved, aseptate or rarely uniseptate, more or
less truncate at both ends, aggregated in a slimy mass.

Chlamydospores often developed on MYA, OA and PDA,
terminal, dark olive-brown, ellipsoid or pyriform, smooth,
thick-walled, 10-15 x 6-10 um, aseptate or rarely unisep-
ate’.

NG Gye NOmOLrOWtl sOCCULS.

Specimen examined: an isolate from air, at Kano-cho,
Ikuta-ku, Kobe-shi, Japan, June 12th, 1980, N. Toyazaki,
NHL 2891, holotype.

Ofechesspecies, currently, assigned sto the genus, C-
dimorpha somewhat resembles C. fertilis Hughes § Kendrick
(1968) because of the crowded appearance of polyphialides
which are produced at the portion where cylindrical coni-
diophores proliferate, and because it forms prominent
phialide collarettes, as well as fusiform, curved and setu-
late conidia. It differs from C. fertilis in that it lacks
sterile setae and has larger, 3-septate conidia and mono-
phialidic conidiogenous cells.

Chloridium codinaeoides (loc. cit.) may be confused
with the monophialidic fruiting structures of the present
Species, but in the former, conidia are smaller 5-9 x
1-1.5 um as opposed to 10-18 x 1-2 um in C. dimorpha.

The another superficially similar hyphomycete, Hypho-
discosia europaea Hol.-Jech. § Borowska has recently been
described by Holubov4-Jechova and Borowska (1981). Both
are characterized by cylindrical, setulate phragmospores
produced acropleurogenously at the elongating part of sim-
ple, erect, pigmented conidiophores. Hyphodiscosia euro-
paea is distinct from the present fungus in that the coni-
dia are sympodial-polyblastic and dark-colored at maturity,
and in lacking a monophialidic conidia.

ACKNOWLEDGMENT

456

We are grateful to Dr. Walter Gams, CBS, for his crit-
ical review of the manuscript and for many helpful sugges-
Cions.

REFERENCES

Ellis Ma Bl oy Ceeeeiore Demat vaccousmily DROMy Cele Samm Dr.
472-474. Commonwealth Mycological Institute, Kew,
England.

Gams, W., and V. Holubova-Jechova. 1976. Chloridium and
some other dematiaceous Hyphomycetes growing on decay-
TN WOO tame CUCTEBMY GO Lramm Ls mrp

Godeas,/ A. M:.) S.G. Marchand @andsD. Gabraly 01977) lora
criptogaémica de Tierra del Fuego. Tome X. Orden Hy-
phomycetales. Fasciculo 1. Fundacién para la Educa-
cién, la Ciencia y la Cultura, Buenos Aires, Argen-
CinavemLloep.

Holubova-Jechova, V., and A. Borowska. 1981. Hyphodisco-
sia europaea, a new species of lignicolous Hypho-
MY.CECES es GESkauMy iO lems ome 2 0 5 lee

Hughes, S. J., and W. B. Kendrick. 1968. New Zealand
fungi 12. Menispora, Codinaea, Menisporopsis. N. Z.
ULB O Ghat Oc meg 25-150, oe

Kornerup, A., and J. H. Wanscher. 1978. Methuen Handbook
ote Colours 5 rdeedsm nme Met huchemLOndClhwmmE > cme

Matsushima, T. 1971. Microfungi of the Solomon Islands
and Papua-New Guinea (Kobe). p. 14.

Matsushima, T. 1975. Icones Microfungorum a Matsushima
Uectorumm (Kobe) Meme ppresOec7e

Morgan-Jones, G., and E. G. Ingram. 1976. Notes on Hypho-
mycetes XV. Two new species of Codinaea. Mycotaxon
4: 504-509.

Pirozynski, K. A. 1972. »Microfungi of Tanzania I. Miscel-
laneous fungi on oil palm. II. New Hyphomycetes. CMI
My,.cO Seb ap ae ZrO LU.

PirozynsSkae Kell rand > 0 ba tds Lo (Ore SOmems etose
Hyphomycetes of leaf litter in South India. Can. J.
BO Cai Gime0 foe

Reddy, S. M., and S. S. Reddy. 1977(1978). A new species
of Codinaea. Sydowia 30: 186-188.

Samuels, G. J., and E. Muller. 1978. Life-history studies
of Brazilian Ascomycetes 1. Two new genera of the
Sphaeriaceae having, respectively, Sporoschisma-1like
and Codinaea anamorphs. Sydowia 31: 126-136.

Sutton, B. C., and C. S. Hodges. 1975. Eucalyptus micro-
fungi: Codinaea and Zanclospora species from Brazil.
Nova Hedwigia 26: 517-525.

MYCOTAXON

eee
p Ole Di Nos 63) pp 45/7 =464 October-December 1981

fhemhsilopezvordatuneieeVvadle
Additions to the genus Pachyella.

Donald H. Pfister

Farlow Reference Library
and Herbarium of Cryptogamic Botany
Harvard University, Cambridge, MA

and

Frangoise Candoussau

22, rue H6o-Paris, 64000 Pau, France

SUMMARY

A revtew of the genus Pachyella ts gtven tn vtew of
recent collections from France and continued study by the
sentor author. One new spectes ts proposed, P. peltata,
and one new combtnatton tn made, P. pseudosuccosa. A key
ts gtven to the nine spectes now tneluded in the genus.

The senior author has revised the genus Pachyella
(Pfister 1973) and emended its circumscription. The
junior author has made significant new collections of
species Of the genus in France. This paper presents one
Mewespeclcsnotm acnyella trom trance, asreportsot.?.
punettspora from France, and a revision of the nomencla-
ture of one species. A revised key to the genus Pach-
yella is given.

The literary history of the genus Pachyella was re-
viewed by =riister (1975). “Sance that Cimesseveral
smaller contributions have been made (Pfister 1974, 1975).
Pristens(1979) also added’ on species to thesgenus er.
hydrophtlta (Peck ex Sacc.) Pfister, and Berthet and Dona-
dini (1978) described Peztza aquatilis which was later
transferred to Pachyella by Donadini (1978). This
species is discussed below.

A recent paper by Donadini (1980) should be men-
tioned. In it Donadini reduced Pachyella to the rank of
a subgenus under Pezgtzga. There is no indication that

458

specimens were studied. Unfortunately several bibliogra-
phical improprieties have occurred.

We feel that Pachyella should be maintained as a
distinct genus. The species are not only morphologically
and anatomically distinct from Pegtza but they are ecolo-
gically quite different as was pointed out earlier by
Pfister (1973).

The following is a synopsis of the species now
placed in Pachyella. Complete synonymies may be found
for previously treated species in Pfister (1973).

SVNODS smo les peeics

Pachyella adnata (Berk. & Curt.) Pfist., Canadwnum pou.

Sle 20 L0te o/s.

Earlier Aleurta annamttica Pat. was added to the
synonymy (Pfister 1974). Few additional collections have
come to light. The distribution is wide but disjunct.
Additional collections deposited in FH have been examined
from Ohio (W. B. Cooke 48304) and Connecticut (DHP). The
species is unreported in Europe. Donadini (1980) listed
the authors as "(Berk. et Currey)" [sic].

Pachyella aquatilis (Berthet & Donadini) Donadini, see P.
pseudosuccosa.

Paehyella babitngtontt (Berk. & Br.) Boud., Hist. Class.
DUSCOM ee LU lea Dime orl) ie
Peztza extditformis Berk. & Br. may be added to the
already lengthy synonymy. Donadini (1980) listed the
authorseor PD. ipabirgton ease (Berke eh Bress)iue| sll.

Pachyella elypeata (Schw.) Le Gale Prodr.
Madagascar’ 4:)277 1953.

This species is still known only from North America

though the new species de
: scribed below from F 1
quite simitar® a Noe

Flore Mycol.

Pachyella hydrophila (Peck ex Sacc.) Pfister, Mycotaxon
8335.) 19798
This species and Pachyella punetispora are quite
Similar anatomically. Detailed field studies of P.
hydrophtla should be made--it is thus far only known from
dried material.

459

Pachyella megalosperma (Le Gal) Pfist., Canad. J. Bot.
ime OL Ieee O73

No recent collections of this species have come to
our attention. Comments on Pachyella pseudosuccosa
should be seen below.

Figure 1. Pachyella peltata. Apothecia from the type
collection. Scale = 2 cm.

Pachyella peltata Pfister §& F. Candoussau sp. nov.

Figures 1 and 2

Apothecta 4-7 em diam, planum, sessilta planae, sim-
tits Pachyella clypeata. Hymentwn brunneum. Exctpulum
extertum ex hyphts 55 x 25 wm composttum. Hyphis exteris
rectts vel undulatts vel ramtficans. Asct J+, 500 x 20-
22 um dtam. Ascosport elltpsoidet, laevt biguttulati
27-350 x 18-15 ym. Paraphyses rectae, brumnnae.

Holotype: sur tronc de Saltx pourri, 4 terre, endroit
inondé périodiquement, avec Pstlopezta nummtlartalts
Pfist. §& Candoussau et Climacodon pulcherrtmus (Berk. §&
Curt.) Nikol. ainsi que sur Alnus gluttnosa au méme
endroit. 14. IX. 1980. Forét de Bugangue-Oloron, Pyré-
nées Atlantiques FH. Isotypes CUP and herb. Candoussau.

460

SSS eon
See

Figure 2. Pachyella peltata. Outer excipulun.
Scale = 50 pm.

Apothecia gregarious 4 to 7 cm diam, sessile, flat,
toward the center undulating--plicate as far as the margin.
External surface dirty-white. Hymenium brown (chatain),
shining, slightly wiscous hlesi se dirtyewa tesUupn ton camm
thick at the center, tinted yellowish where broken. In
external features agreeing closely with P. elypeata
(Schw) theaGale

Outer excipulum of longitudinally elongate cells
with their long axes oriented perpendicularly to the
outer Psurfacero ts theceapotiecium ao oece ll Smlecp acc
larger of the *célissmeasuringe ome co impor es li ehie by
Jarger..* Cells tichtly *packedeandvonl\es lish tlyecwollen
in width. The outer cells are uniformly embedded in a
gelatinous matrix and give rise to hyphoid hairs of 5 to
7 ym in width to more than 400 pm in length, which often
branch and anastomose or become moderately coiled.

Medullary excipulum of dense texturasintricata ce!
sparingly present, hyphae 6-12 um in diam.

Asci diffusely J+, nearly 500 pm long and 20-22 pm
broad. :

Ascospores at maturity hyaline, thin-walled, smooth
or with some cytoplasmic granulation, biguttulate but
often jobscurel yrsone 2/50 exe Solo anim

Paraphyses septate, enlarged above, exceeding the
asci, of granular contents which become dark brown in
freshamatenivaluwheneplacedmins1odine.

This species is quite similar to Pachyella clypeata;
the spores are larger and the configuration of the

hyphoid hairs differszaasinjhabital and erowth forme tars
pdentical.;

461

Figure 3. Pachyeila punetrspora. Apothecia from
material collected in France. Scale = 1 cn.

Pachyveugapunertspora Prist, , Canad. Js Bot. ol: 2019.
SR Figure 3
= Pezitza puncttspora (Pfister) Donadini. Doc. myco-

logiques 11: 26. 1980.

Pierre Gabard, Guy Roux and the junior author have
collected this species in France (Pont du Goa, Vallée du
poussucou, 900 m alt. 15.VII.1979; Crampes de Piet,

Pallee d!'Ossau. s00 malt. Leg. Guy Roux, Pyr. Atlan-

tiques, environs de Pau, s/Betula et autres bois, 28.VI1.

1979). These collections constitute the first report of

the species outside North America. Figure 3 shows apothe-

cia of the French collection.

Pachyella pseudosuccosa (Le Gal) Pfister, comb. nov.
Galaetinta pseudosuccosa Le Gal, Rev. Mycol.
(Paris) 10: 91-95. 1945.
Pegiza aquatilis Berthet & J.-C. Donadini. Bull.
Mens. Soc. Linn. Lyon 47: 205. 1978.
Pachyella aquitilts (Berthet & Donadini) Donadini,
BuliSoce Linn. Provence 317.518251973,

When Le Gal described Galactinta pseudosuccosa, she
did so in such a way as to introduce some ambiguities,

Dee)
ll

462

particularly in clearly stating the name in association
with the Latin description. It seems to be validly
published. One form she described, forma macrospora, was
the form upon which Galactinia megalosperma (Le Gal 1953)
was based. The typical form, material of which has been
unavailable for study, was described from dried material
in the Boudier herbarium and was collected by Barla in
NIC Cw iim ode

A portion of the holotype specimen of Pegtza aquatt-
its, kindly sent by M. Berthet wapnees Closclyswatiimtnat
described by Le Gal as Galactinta pseudosuccosa. Both
have a reddish brown hymenium and both are said to
produce a yellow juice when broken. Moreover, the Barla
collection and those of Donadini are all from the same
general area in S. W. France:

Pachyella vtolaceontgra (Rehm) Pfist., Canad. J. Bot.

aig aval absyats

Previously the combination of this species in Galac-
LENCGaDY sBOUdL Cy (HUStyeG1asS mp Us COM mc Liem) m4 eel OU @)
was overlooked. Also to be added to the synonymy is the
combination: Pegziza vtolaceontgra (Rehm) Smits'Ka (Ret-
SLUSOVIeGri Di Ukrayiniod KLCV) ep en oeLo >) mm onadi nd
(1980) also proposed the combination Peztza vitolaceontgra.

The junior author has made collections of Pachyella
vtolaceontgra in France (Bois de St. Maur-Mirande, Gers,
13.VIII.1971; Forét de Bugangue, Oloron, Pyr. Atlantiques
Do ee A975 HH HOLD ee Reeeb . ekhOF Geen el mt Ga NGOUSS ALI.
Other collections were made in Switzerland by A. Nyffe-
negger (near Kanton Bern: FHjZ)" and by iBruno Erb and
Stappung Erlinsback (automne 1980, Herb. F. Candoussau).

KevVeCOms DCClCSmO LE UaiiC11 a
Hyphoid hairs forming a aca pean at the base of

Giicma DOC Ne Cl aa sun ue A cea a, ey
Hyphoid hairs not Revandlays a Hsccultayo Palisade layer
aimee NemDaAsceOlmenceapOthectam.: 2). oll

2. Ascospores smooth 13-16 x 18- 25 (28) “pam. “Aieerdacosia
UMDeRecOMChestiut color,.outer layer of exci pulum
of radially arranged sileinage célls, hyphoid hairs
neither branched nor erat ae N. America.

: ve: fyi heels Ribena

Ty Aecospannes or r apothecial aonstaneenan not as
ADOVC Wms. 5

Ascospores snensal 13- 15 x 27- 30 si, ayterd ice ihe dese

HMDeymOuL CG slay eCTsOLmexcapulumeot strontly packeds

only slightly swollen cells, hyphoid hairs branched

Wiel AWB Sie Wi. ONS. 6 ap 66 0 0 6 Wee jeune:

nS COSDOLeSaVanlOUs Ly sOrlamenucdm., |.) eunsn po the cn ta

4, Ornamentations taking the form of iene eiheve
topped warts, N. America, West Indies, and Japan.

spat ean eC OnOLa
A’ rene nar anes Ss ikiise in ane aovatl of warts or
I MECTACO MNO EM NY LEMS 4 «6 Se Ae

Spores ornamented with anaatetostins fi hemartiem
reddish brown, known aes from Madagascar .
: rae RRC LRECAE
Sneiwes ermented rene agit CEeReS Bo 4 (ho ee Te
6. Hymenium dark brick to sepia, when damaged not
producing a yellow ‘aig Busopemr
: fee Se apererord sk
6. abareneiwi aati ovat nen damaged producing a
velloW julce, burope’. 9... « .« 4. pseudosuccosa

Spores smooth or punctate, margin attached to the
substrate, apothecia Soon NO te Siea vet mtnanmt ac

in diam, worldwide i ce Ney her Meme. DOD neLOnTe
Spores ornamented with Se eens, margin free, apo-
Checram ccictal |yelarcernsthanwl acm. mae 8

Sem opores el 2—1 55x. 21-25, 71m, marked neice smell occ
discernable warts, el up to 9 pm, N.
ANEGGECa, sEUrOpe wm. . .P. punettspora

Seem porese | S=10exe22— 26 um, marked with warts which
can, scarcely be seen with the light microscope,
satis Ups tOmlo pithinadial iN. eAMCTC acme cme

: Pe Ae ee oy ee Bo. tac hydrophila

463

464

ACKNOWLEDGEMENTS

We wish to thank Richard P. Korf for his comments on
this manuscript, seeror siberchets Kind lyescitemaveria imo t
Peziza aquatilts for our study. We are grateful to Emile
Jarias and Guy Roux. tor photographs. slhe senior author's
work was supported in part by National Science Foundation
Grant sDEBPs02340135-

DL PERATURES CITED

Berthet, P. and J.-C. Donadinuss l97o8 Un nouveaurlezrca
Dy ex, Oe AManSe "lest cop dguatT Tom nOV eS) amebU aie
Mens. Soc. Linn. Lyon 47: 202-205.

Donadiniy J0-C8 919/87 a eLesyenresrez? sae epcimodl Nt
Amans (II). Les Pezizes de Haute-Provence et de
Dauphine-Savoie. Bull. Soc. Linn. Provence 31: 9-36.

1980. Le genre Peztaza (Dill.) Linne per
Saint Amans sous-genre Pachyella (Boudier) nov.
comb. Doc. mycologiques. 11: 25-26.

Bilstein el pie OLS ees DSMLODeZLOLGm LUN Minmmmn VemmiC

genus Pachye Lite Canad Jam ROU Od ez COG cUeoe
~ 819742) Ibid Vl Alew-taraunane bi care

synonym of Pachyella adnata. Mycologia 67: 181.

1975s o€anningnel ectronemicrographcmores
ascospores of Pachyella (Discomycetes). Mycotaxon
3: 105-108.

1979. Type studies in the genus Peztza
VI. Species described by C. H. Peck. Mycotaxon 8:
333-338.

MY COTA XON

a Ee
VOLS XIII," No. 3, pp. 465-468 October-December 1981
en ee ee ee pr ee Ra ee

PITHOMYCES PAVGII, A NEW COMBINATION FOR
TRICHOCLADIUM PAVGII AND PITHOMYCES FUNICULOSA

MARY E. PALM and ELWIN L. STEWART

Department of Plant Pathology
Universtty of Minnesota
SE Louse MN o0lUS USA

Trtchocladtum pavgtt Nath was published as a new spe-
cies from India on 16 July 1979 (1). Additionally, a new
dematiaceous hyphomycete from Swaziland was described as
Pithomyces funiculosa Palm, Stewart, & Rossman on 19 Octo-
ber 1979 (2). We have determined that 7. pavgti and P.
funteulosa are conspecific. Trichoeladtum pavgit has pri-
ority according to the International Code of Botanical
Nomenclature, Article 11.3 (3). However, based on conidium
development and secession, 7. pavgtt can only be accommo-
dated in Pithomyces Berk. & Br. Trichocladium Harz has
been broadly circumscribed and at present contains a hetero-
geneous assemblage of species which produce transversely
septate conidia holoblastically. Pithomyces, in contrast,
is characterized by the holoblastic production of dema-
tiaceous conidia which are usually both transversely and
obliquely septate. Moreover, rhexolytic conidium seces-
sion leaves a characteristic denticle on the conidiogenous
cell. Trichoeladium pavgit must be transferred to
Pithomyces. The new combination is herein made and the
taxon redescribed based on additional studies of isolates
from India and the United States.

Pithomyces pavgit (Nath) Palm, Stewart, & Rossman, comb.
nov. Figs. 1-4
= Trichoeladium pavgit Nath, Sydowia 31: 122-125. 1978.
(basionym)
= Ptthomyces funiculosa Palm, Stewart, & Rossman,
Mycotaxon 10: 246-250. 1979.

COLONIES on malt agar white to grey becoming brown to
black, floccose to appressed, dark ropy strands of conidio-

466

phores frequently present. HYPHAE septate, branched, of two
types, hyaline to subhyaline, smooth, 1.5-5 (x=2.8) um diam,
and subhyaline to pale brown, sometimes finely roughened,
2.5-6 (x=3.4) um diam. CONIDIOPHORES micronematous to semi-
macronematous, mononematous, frequently funiculose, infre-
quently branched, septate, subhyaline to pale brown.
CONIDIOGENOUS CELLS holoblastic, monoblastic or polyblastic,
integrated, intercalary, determinate or seemingly prolifer-
ating percurrently to produce one or more conidia, cylin-
dric to doliform, denticulate following secession of
conidia, denticles short cylindric to doliform, 1-4 (x=2.4)
xX 1.5-4 (x=2.3) um. CONIDIA solitary, pleurogenous,

mostly verrucose, pale to dark brown, clavate to obovate

to obpyriform to broadly or fusiform ellipsoid to reniform,
transversely (0-)1-3(-4)-septate, mostly 2-septate, may

be slightly constricted at the septa, rarely with 1-2 ob-
lique or longitudinal septa, 10-17 (x=13.3) x 4.5-12

(x=7.4) um (1l-septate), 13-20 (x=17.2) x 5-12 (x=7.8) um
(2-septate), 17-26 (x=22.1) x 6-12 (x=8.3) um (3-septate) ,
bases 2-4 (x=2.6) um diam, with a portion of the conidio-
genous cell usually remaining as a basal frill.

Collections examined: India: Hyderabad, Rajendranagar,

V. Ravindra Nath. From kernels of Arachts hypogaea L.

3 i 1973 (IMI 205718 [holotype] and living culture received
from V.R.N.). Swaziland: Luyengo, A.Y. Rossman. From .
fungal comb of Macrotermes ukuzit Fuller. 15 1x 1977
(MPPD). United States: Alabama, Chambers Co., G. Morgan-
Jones. From municipal industrial waste-water. v 1980
(AUAM 2405).

In addition to its occurrence in India and Swaziland,
G. Morgan-Jones (personal communication) has isolated P.
pavgtt in the United States. Based on our observations
using phase contrast and scanning electron microscopy, the
conidiogenous cell either proliferates percurrently through
the denticle, which remains as a short annellation, or the
apparently inelastic outer wall ruptures and remains at the
base of the exposed, elastic inner wall. Remnants of
conidiogenous cells surrounding the possibly proliferating
conidiogenous cell are seen infrequently and mainly in
Older cultures. Ravindra Nath's line drawings of P. pavgtt
indicate flared structures resembling annellations at the
apex of some elongate conidiogenous cells which he de-
scribed as "roughened". Examination of the type material
(IMI 205718) and of a living culture indicate that the

467

FIGS. 1-4. Ptthomyces pavgit. 1. Photomicrograph (phase
contrast microscopy for Figs. | and 2) of a developing co-
nidium with intact outer wall. X2000. 2. Photomicrograph
of a conidiogenous cell following rhexolytic conidium se-
cession showing remnants of either the inner and outer walls
or the walls from two successively produced conidia. X2000.
Figs. 3-4. Scanning electron micrographs of conidium and
conidiogenous cell with inelastic outer wall beginning to
rupture (Fig. 3) and conidium and conidiogenous cell with
ruptured outer wall or a possible annellation (Rid. 4s
X3000.

468

fungus sometimes appears to proliferate percurrently, but,
a conidiogenous cell so long and with more than one
"annellation" was never observed by us. Final elucidation
of conidiogenesis requires ultrastructural or time lapse
Studies or both.

ACKNOWLEDGEMENTS

This work was made possible in part by funds provided
by the USDA, Forest Service, Forest Products Division
(Nos. 12-39 and 12-144). We thank Dr. G. Morgan-Jones for
the opportunity to observe and cite his isolate of P.
pavgtt and appreciate the helpful comments and suggestions
of Drs. G. Morgan-Jones and W.J. Sundberg regarding the
manuscript. Minnesota Agricultural Experiment Station,
Scientific Journal Series, Paper No. 11,777.

LITERATURE CITED

1. Nath, V. Ravindra. 1978. On Trtchoeladium and a new
species, Trtchocladtum pavgtt. Sydowia 31: 122-125.

ac) let Piy Wikteon pbs Sacaicteien cule ais MOSSE I
Pithomyces funiculosa sp. nov. from fungal comb of
Macrotermes ukugtt in Swaziland. Mycotaxon 10:
246-250.

Sous Catlett Ases Ct a t0eds 1) melo / Cement erat Orde
Code of Botanical Nomenclature adopted by the
Twelfth International Botanical Congress, Leningrad,
July 1975. Regnum vegetabile 97: 10.

MYCOTAXON

VOU HEL NOS spp.) 469-4:7.0 October-December 1981

VALIDATION OF THE HERPOMYCETINEAE AND
HERPOMYCETACEAE IN THE LABOULBENIALES

ISABELLE |. TAVARES

Department of Botany, University of
California, Berkeley, California 94720

When the Euceratomycetaceae in the suborder
Laboulbeniineae was described (Tavares, 1980), a comparison
was made between the development of the perithecia in the
Laboulbeniineae and the Herpomycetineae I. Tav. (nom. nud.).
Inadvertently, I omitted a reference to the second volume
of Thaxter's monograph (1908) of the Laboulbeniales, in
which the tribe Herpomyceteae was described in the key to
genera within the family Laboulbeniaceae. In addition,
there was no mention of the family Herpomycetaceae, nor was
a diagnosis of the Herpomycetineae included (Tavares, 1980).
Consequently, a Latin diagnosis for the Herpomycetineae
follows, for which I am indebted to Dr. William J. Dress,
Bailey Hortorium, Cornell University.

Subordo Herpomycetineae nom. nov. Dioeciae. Thallorum
femineorum perithecia uno vel pluribus axibus receptacular-
ibus secondariis, qui una vel pluribus cellulis suprabasal-
POUS@ileaxeupLimardiouproducti sunt, porntata. —Peritheci7
paries exterior ex seriebus quattuor verticalibus cellular-
um constans, serie quaque ex cellulis numerosis brevibus
altitudine subaequis composita. Cellula infima in una
serie verticali cellularum parietalium perithecii juvenalis
et introrsum et sursum per medium perithecii crescens,
carpogonium formans. Antheridia sunt phialides.

Familia Herpomycetaceae (Thaxter) stat. nov.
Tribus Herpomyceteae Thaxter. Mem. Amer. Acad.
Weta iesig TE ABW AER They

Family Herpomycetaceae: As in the suborder.
Dioecious, occurring on cockroaches (Blattaria). The ger-
minating spore becomes transformed into a primary axis of
four or more cells. In the female, perithecia are produced

470

on secondary axes arising from the primary axis. Vertical
septa are formed in the perithecial primordium, producing
four cells, each of which divides into a vertical row of
wall cells. When there are two tiers of wall cells, one of
the four lower cells grows inward and upward and forms a
central carpogonial upgrowth (Tavares, 1980). At maturity,
there are many short outer wall cells, equal or subequal in
height, in each vertical row. Four vertical rows of inner
wall cells surround the passage in the perithecial neck
through which the ascospores emerge (Tavares, 1966). Eight
spores having median septa are formed in each ascus. Male
thalli bear elongate, simple antheridia.

Thaxter's description (1908) of Herpomyceteae, which
was only a brief statement in his generic key, neverthe-
less served satisfactorily to distinguish the tribe, be-
cause Herpomyces, the solitary genus, is the only dioecious
genus having secondary receptacles and simple antheridia.
Although Dimeromyces aberrans Thaxter (1924) may have
simple antheridia, additional receptacular cells in the
female thallus are produced in the primary axis and do not
form a secondary axis as they do in the closely related
genus Dimorphomyces, in which all antheridia are compound.

In his poorly preserved material, Thaxter was unable
to see the internal structure of the perithecium of
Herpomyces (see Thaxter, 1931). Consequently, he (Thaxter,
1908) placed Herpomyces in the Laboulbeniaceae because of
the simple antheridia, not realizing how strongly its peri-
thecial development differed from that of the other genera
invthis family.

LITERATURE CITED

Tavares, Isabelle I. 1966. Structure and development of
Herpomyces stylopygae (Laboulbeniales). Amer. J. Bot.
D3) BLI=3 18.

- 1980. Notes on perithecial develop-
ment in the Euceratomycetaceae fam. nov. (Laboulben-
iales, Laboulbeniineae) and Herpomyces (Herpomycet-
ineae). Mycotaxon 11: 485-492.

Thaxter, R. 1908. ) Contribution toward, a monograph of the

Laboulbeniaceae. Part II. Mem. Amer. Acad. Arts Sci.
133 217-4693. Pls. Px LP xXx,

- 1924, Contribution towards a monograph of the
Laboulbeniaceae, Part III. Mem. Amer. Acad. Arts
Soin Pi 2309-406 2 ais elle

« L931. “thrid Ss Part Ve" Mem. ‘Amer. Acad. vArts
SCi ty elGst 1 —43 570 ee bs el bo

MY COTAXON

VON ELS eNO. 3, pp. 4/1=480 October-December 1981

ERYNIA (ZYGOMYCETES: ENTOMOPHTHORALES ) :

VALIDATIONS AND NEW SPECIES

RICHARD A. HUMBER

USDA-SEA-AR Insect Pathology Research Untt
Boyce Thompson Institute at Cornell
HOUe TRO MECHA CUMmmuNeCW VOR K EL GO, OSA

SUMMARY

New specific names are proposed to validate three
heretofore invalidly published species of Hrynta
Nowakowski (Entomophthoraceae): £&. bullata Thax-
ter & MacLeod in Humber, #. creatonott Yen in Hum-
ber, and £. myrmecophaga Turian & Wuest in Humber.
Erynta formtcae Humber & Batazy in Humber, sp.
nov., iS proposed as a name for a fungus occurring
on ants and differing from the incompletely char-
acterized #. myrmecophaga.

During recent studies on the entomopathogenic Entomoph-
thorales, Hrynta Nowakowski and the closely allied or syno-
nymous genus (depending on one's taxonomic opinion) Zooph-
thora Batko have received much attention (Remaudiére and
Hennebert 1980; Remaudiére and Keller 1980; Humber 1981,
1982; Humber and Ben-Ze'ev 1981; Ben-Ze'ev and Kenneth 1981,
1982). Several species referable to Hrynta sensu Humber &
Be-Ze'ev (1981) are not validly published under the Inter-
national Code of Botanical Nomenclature. This paper vali-

472

dates three of these species and proposes a second new spe-
cies of Erynta from ants.

ERYNIA BULLATA Thaxter & MacLeod, sp. nov.

CORPORA HYPHALIA vartabtlts per tntttum contagtonts, hyphotdea postrem.
NUCLEI persaepe consptcut, mtntmum 6-10 um tn dtametro, contentts gra-
nulatts autem stne nucleolo consptcuo ovotdeo centralt. CONIDIOPHORA
dichotome digttate ramosa ad apices, intertexta tn hymentum continuum;
tn taentts emergens et itn integumenta super corpore hospttts coalescens.
CONIDIA PRIMARIA untnucleata, bttuntcata, elltpsotdea ovotdea, 23.0-87.5
wn x2 12.0-16.5 wm (medto 29.5 x 14.0 um), expulsa vehementer everstone
paptllae distinetae contra contdtophora. CONIDIA SECUNDARIA tnobser-
vata. SPORAE PERDURANTES hyalinae vel stramineae vel palltdae alutaceae
tn massa tn corpore hospttis; multtnucleata ubt maturae; eptsporits con-
spteue vel bullatts vel tuberculatis, 37-62 \wm (medto 50 um) tn dtametro
toto bullts tnelusts. RHIZOIDEA numerosa, stmplicta, hyphotdea, stne
haptero disctform terminalt, ex abdomine et thorace emergentta; in
praesentta vel conidtorum vel sporarum perdurantium formata. CYSTIDIA
desunt.

HYPHAL BODIES variable in size and shape during early stages of develop-
ment in host, becoming hypha-like at time of the host's death. NUCLEI
easily distinguished in most Vegetative or reproductive structures;
large, usually 6-10 um or more in diameter; contents granular, without
a prominent central ovoid nucleolus. CONIDIOPHORES branched digitately
toward the apices, intertwining to form a continuous hymenium; emerging
in bands from weak points of the host exoskeleton, coalescing to form
more or less complete coverings over the abdomen and the junction of
the neck and thorax. PRIMARY CONIDIA uninucleate, bitunicate, ellips-
oid to ovoid, 23.0-37.5 um x 12.0-16.5 um (averaging 29.5 x 14.0 um),
forcibly ejected by the eversion of the conspicuous basal papilla
against the conidiophore. SECONDARY CONIDIA not observed. RESTING
SPORES forming a compact, hyaline to creamy-yellow or light tan mass
inside the host body; multinucleate when mature; individual spores

with prominent, rounded bullations up to 5 Lm in height on the epi-
spore; 37-62 um (averaging 50 um) in overall diameter (including the
episporial bullations). RHIZOIDS numerous, unbranched, hypha-like,
without any terminal disc-like holdfast, emerging principally from the
abdomen and thorax; present whether the fungus produces conidia or
resting spores.

HOLOTYPE: CUP 58970 (Cornell Univ., Plant Pathology Herbarium).
Resting spores in Sarcophaga aldricht. Coll.: D. Tyrrell,
235 June 1981.

PARATYPE: CUP 58971 (Cornell Univ., Plant Pathology Herbarium).
Conidia on Sarcophaga aldricht. Coll.: D. Tyrrell, 23
June 1981.

TYPE HOST: Sarecophaga aldricht Parker (Diptera: Sarcophagidae).
TYPE LOCALITY: Birch Creek Road, Webwood, Ontario, Canada.

473

This species appears to be a relatively common patho-
gen of calliphorid and sarcophagid flies in North America.
It was collected several times in the northeastern United
States by Roland Thaxter (Riddle 1906, as Entomophthora "x")
but never formally described. After Thaxter's death, Povah
(1935) received permission from Dr. W. Weston at Harvard
University to use Thaxter's unpublished herbarium name,
Entomophthora bullata, for the fungus Povah found attacking
bluebottle flies in Michigan. This name, attributed to
Thaxter, was presented with scant details of the fungus's
morphology except for the bullate nature of its resting
spores (33-50 um diam. in Povah's collection) and the over-
all similarity of the primary conidia of this fungus with
those of Erynta amertcana (Thaxter) Remaudiére § Hennebert
(1980). Unfortunately, Entomophthora bullata Thaxter in
Povah appeared in print after 1 January 1935 and is, accor-
ding to Article 36 of the International Code of Botanical
Nomenclature, invalidly published without a Latin diagnosis
or reference to any previously published Latin diagnosis.
All subsequent combinations based on this name — JZooph-
thora bullata (Thax. in Povah) Batko (1966) and Hrynta bul-
lata (Thax. in Povah) Remaudiére § Hennebert (1980) — are
also invalidly published.

The most thorough study of £. bullata is that by Mac-
Leod et al. (1973) who provide a complete morphological
characterization. They suggest the host's age may affect
WiectjerecOnralasor resting’ Spores. are produced at the cime
of the fly's death. Kramer (1979) studied the time course
Simin tectionsandstne spore states sprodcuccdsin whreesspectes
Dimcagierpiottdmrt les mintcctedein tno =lavOraloLy se sicmCcOLLO-
borated that the disease in younger flies does tend to pro-
duce only conidia while the probability for resting spore
production increases with the age of the flies at the time
omeintecr one minecontrast to MacLeod et al (19/5), uwiic
state that the fungus never produced both conidia and
resting spores in the same cadavers of field-infected S.
aldricht, Kramer (1979) found that, on rare occasions, both
spore states may occur together in affected individuals of
Phormia regina (Meig.) and Phaenicta sertcata (Meig.). It
is significant, then, to note that in the field-collected
examples of Sarcophaga aldrtcht in the holotype collection,
tufts of conidiophores are apparent at the joints of the
legs whereas the fly bodies are filled with resting spores.
No bands of conidiophores are apparent on the abdomens of
flies in the holotype collection.

474

Cytological studies by Riddle (1906) indicated that
three-month-old resting spores of #. bullata are multinu-
cleate. This condition was verified with resting spores
from the holotype collectionestained wictieeci1 tncrsaccto-
orcein or bismarck brown. Humber (198la) discusses the
implications of this multinucleate condition in entomoph-
thoralean resting spores.

ERYNIA CREATONOTI Yen, sp. nov.

"CONIDIA pyrtformia, uninucleata, med, 32 x 25 wm, bast paptllata, muro
tenut, cytoplasmatae granoso; NUCLEI med. 12 x 9 \1m; CONIDIOPHORI ramost
coalescentes tegitmenta suffusco vel roseo super totum hospitis corpus.
CORPORA HYPHALIA brevia, segmentis hypharum tnecomposttis, 8-28 ym latis
20-24 ym Longts. CYSTIDIA raro tnventuntur. SPORAE PERDURANTES azygo-
sporae (zygosporae pauca tnventae), globosae; eptsporto laevt atque
crasso; cytoplasma granosum vel vacuolatus; dtam. 10-28 um. Hospes
substrato RHIZOIDEIS ftxus." (Yen 1962)

TYPE: Latin and English descriptions and figure 1-4, J. Insect Pathol.
4, 88-94 (1962).

TYPE HOST: Creatonotus gangis (L.) (Lepidoptera: Arctiidae)
TYPE LOCALITY: Kun-Kuan, Taipei, Taiwan.

This species, originally proposed with the orthographi-
cally incorrect name Entomophthora creatonotus Yen (1962),
was invalidly published since no type was designated. The
combinations Zoophthora creatonott (Yen) Batko (1964) and
Erynta ecreatonott (Yen) Remaudiére § Hennebert (1980) are
also, thererorem invalid! ye pubiisned mas! 1emCchOlCesO mete
description and figures of #. creatonott to typify this spe-
cies is motivated by the lack of any mention by Yen (1962)
of the permanent deposition of either any specimens or the
Slides from which his photomicrographs were prepared. The
English description of all morphological characters of this
fungus is meticulously detailed even if the figures are of
Tim cCedauts li oye

The placement of this species in Erynta is fully
justified by its uninucleate (presumably bitunicate) coni-
dia borne on digitately brached conidiophores. The large
nuclei are readily detected, even in unstained material
(Yen 1962), and correspond to those of all other Frynta
Species. The presence of fascicles of hypha-like rhizoids
and of a few cystidia supports this generic assignment.

475

ERYNIA FORMICAE Humber & Bazazy, sp. nov.

CORPORA HYPHALIA trregularta per intttum contagtonts, 35-155 um x
8-16(20) wm, et posterius mantfestius hyphotdea et saepe parciter sep-
tata. NUCLEI persaepe conspteut, magnt, contentis granulatis autem
sine nucleolo ovotdeo centralt. CONIDIOPHORA in taentis distinctis
emergentia, dtchotome vel digitate ramosa ad apices. CONIDIA PRIMARIA
unitnucleata, bitunicata, obovata vel pyriformia, 18-25 um x 10-16 um,
expulsa vehementer everstone paptllarum contra conidiophora; paptlla
basalt altquantum distincta. CONIDIA SECUNDARIA contdia primarta stmu-
Lantta quanquam parvtora, 16-19 um « 10.5-13.5 um, expulsa vehementer
tn modum contdiorum prtmartarum. RHIZOIDEA sparsa crassa, haptera
disetformta terminalia, ex junetura promesothoracica in fasciculum
laxum emergentta. CYSTIDIA et SPORAE PERDURANTES tgnotae.

HYPHAL BODIES irregular in length, diameter, and branching pattern in-
side host body during early stages of development 35-155 um x 8-16(20)
um, later becoming infrequently septate and more obviously hyphal in
appearance. NUCLEI conspicuous in most vegetative or reproductive
Structures «sarge, With granular contents but no central nucleolus:
CONIDIOPHORES emerging in distinct bands from weak points on the insect
exoskeleton, dichotomously or digitately branched at the apices. PRI-
MARY CONIDIA uninucleate, bitunicate, obovate to pyriform, 18-25 um x
10-16 um, forcibly discharged by eversion of the more or less distinct
papilla against the conidiophore. SECONDARY CONIDIA like the primary
but smaller, 16-19 um x 10.5-13.5 um, forcibly discharged as in the
primary conidia. RHIZOIDS relatively few in number, stout, terminating
in a discoidal holdfast; emerging from the ventral thorax just behind
the head, forming a loose fascicle. CYSTIDIA may be present; if so,
tapering to a point from a relatively broad base. RESTING SPORES not
observed.

HOLOTYPE: CUP 58975 (Batazy 1285a), coll: J. WiSniewski and A.
SOKOLOWSKin m1 9eSept 91977, slucholaspines forest, Poland

PARATYPES: CUP 58976 (Batazy 1286), CUP 58977 (Batazy 2074a) and
specimens retained by S. Batazy and R. A. Humber.

TYPE HOSTS: Ants of the genus Formtca (Hymenoptera: Formicidae): F.
polyectena and F. rufa.

TYPE LOCALITY: Pine forests near Tuchola, Poland.

A comparison of entomophthoraceous fungi reported to
attack ants in Switzerland (Turian and Wuest 1969, 1977),
the Federal Republic of Germany (Loos-Frank and Zimmermann
1976), and Poland (Batazy and Sokoktowski 1977) indicates
the existence of two distinct species of Frynta. FErynta
formtcae is described for the better characterized fungus
(Loos-Frank and Zimmermann 1976; Batazy and Sokotowski
1977); these collections have been attributed to the inval-

476

idly published and less completely characterized Zoophthora
myrmecophaga Turian & Wuest (which is validated below as
Erynta myrmecophaga Turian §& Wuest).

The conidia of the Swiss fungus are smaller (averaging
18 x 12 um), considerably more ovoid, and have a less con-
spocuous papilla than those of F. formcae. The conidia of
the Swiss fungus but not of #. formcae have a single large
vacuole occupying most of the conidial volume. In addition
to being held onto grass blades by the mandibles and clasped
legs, #. formeae further secures the ant cadavers with a
few stout rhizoids emerging in a loose fascicle from the
ventral thorax (Loos-Frank and Zimmermann 1976; Batazy and
Sokotowski 1977). This sort of rhizoid generally terminates
in a discoid haptor (Batko 1966; Humber 198la); although
Loos-Frank and Zimmermann (1976) refer to a holdfast disc
("Haftscheibe') on the rhizoids, neither they nor Batazy and

Sokotowski (1977) illustrate the morphology of this holdfast.

Erynta myrymecophaga has no such fascicle of rhizoids; Turian
and Wuest (1977) report the presence of a few very fine’ rhi-
zoids emerging from all portions of the host body.

The presence or absence of cystidia in £. formticae re-
mains uncertain. Loos-Frank and Zimmermann (1976) note the
presence Of isolated, broadicystidlantaperingmtordampoiet,
but do not state where these occurred on the host, or at.
what stage of development. Batazy and Sokotowski (1977)
observed no cystidia on any specimens allowed to develop and
sporulate in damp chambers.

Loos-Frank and Zimmermann (1976) provide a detailed
study of the effects of #. formtcae on its host, and find
the behavior of affected ants and fungus-mediated histo-
pathology of the brain to be much like the effects in ants
caused by the parasitic trematode Dicrocoeltum dendrtttcun.
Humber (1975, 1976) also found a marked tendency for hyphae
of Stronguellsea magna’ Humber to penetrate the thoracic gan-
glion and brain of affected flies, but without causing any
obvious histopathology or changes in host behavior.

More complete information about the morphology and
pathobiology of EF. fornteae is provided by Loos-Frank and
Zimmermann (1976) and by Batazy and Sokotowski (1977).

477

ERYNIA MYRMECOPHAGA Turian & Wuest, sp. nov.

CONIDIOPHORA paretter dtchotome ramosa; tn taenits ctreum abdomen inter
abdomen et ocetptttum et etreum partes orts emergens. CONIDIA PRIMARIA
uninucleata, bitunicata, late ovotdea vel pyrtformta, medio 18 um x 12
um, vacuolam centraltam magnam conspicuam et guttulas oleosas sudano-
phtlas numerosas continentia; paptlla basalt indistineta. CONIDIA
SECUNDARIA contdia primaria stmulantta stne vacuola quanquam parviora.
Hospes ad substratum mandibults et crurtbus adhaerens.

CONIDIOPHORES sparingly dichotomously branched, emerging in bands on

the abdomen, between the head and thorax, and around the mouth. PRIMARY
CONIDIA uninucleate, bitunicate, braodly ovoid to pyriform, averaging
18 um x12 um, containing a large central vacuole occupying most of the
conidial volume and numerous sudanophilic oil droplets; basal papilla
indistinct. SECONDARY CONIDIA similar to but smaller than primary
conidia, but not containing the prominent central vacuole. Host at-
tached to the substrate by the mandibles and clasped legs.

HOLOTYPE: Figs. 1, 2, and 4, Mtt. Schwetz. Entomol. Ges. 50:285-
289" 61977)%

TYPE HOST: Servtformica fusca L. (Hymenoptera: Formicidae).
TYPE LOCALITY: Sparse grasslands near Geneva, Switzerland.

This description of £. myrmecophaga is obviously frag-
mentary. There remains enough reasonable doubt about cer-
tain aspects included in the Latin description provided for
Zoophthora myrmecophaga (Turian and Wuest 1977) that it was
preferable to provide a wholly new description including
only those characters which seem relatively well estab-
lished.

Turian and Wuest (1969) found no rhizoids attaching
ants affected by EF. myrmecophaga to the substrate. These
authors later described very fine rhizoids (Turian and
Wuest 1977; Fig. 4) which are little distinguished from the
epidermal hairs on the grass blades to which the ants are
affixed. This characterization may actually note tru rhi-
zoids of E. myrmecophaga, but their appearance is not typi-
cal of other entomophthoralean rhizoids. The possibility
remains that these structures, observed on insects collected
in 1967 and 1968, might have been either strands of insect
or spider silk or the fine hyphae of some saprobic deutero-
mycetous contaminant.

478

Similarly, it is so unlikely that the two different
types of restingespores notedeby. lurianeandeWues te GlJ0 7.
1977) actually represent any over-wintering spores which
might be produced by #. myrmecophaga that these spore types
were intentionally omitted from the description and typifi- |
cation of ‘this fungus vabove. “lurlanvandeWuest 41969 shic:
3) first reported finding one solitary resting spore among
the hyphaly bodiesmin’ onevant;, = this se lobosessporcenased
Single-layered, relatively thin, smooth-surfaced wall, was
20 um in diameter, and contained numerous oil droplets. If |
this was an entomophthoralean resting spore, it was immature |
and neither its size nor structure could be considered to be |
taxonomically significant. Turian and Wuest (1977; Fig. 3)
later found a. few globose resting spores with thick walls |
and rugose surfaces; the illustrated structure does not have |
the thick, two-layered wall which is characteristic of ento-
mophthoralean resting spores. Neither were these rugose
spores found in the numbers or aggregated mass in the abdo-
men that one might expect for naturally produced entomoph-
thoralean resting spores. The best clue to the identity of
these structures comes from the conspicuous furrows on the
surface; this morphology is unprecedented in the Entomoph-
thorales but is strongly reminiscentoot tricolipate polien
grains like those produced by oaks and beeches (Fagaceae)
(Faegri and Iversen 1975; Bassett et al. 1978).

Erynta myrmecophaga obviously requires further study
from both any existing specimens and any future collections.
Important. differences between this species: ands thestarepet—
ter characterized #. formtcae exist in the size and shape
of the primary conidia, and the presence and nature of rhi-
zoids. It should be noted, however, that the behavior and
appearance of affected ants, the; relatively low desree of
conidiophore branching, absence of cystidia, and absence of
verifiable resting spores are common characteristics of
these species. s Noneroreche simmanrtulesaromsuttucten til valine
portant or convincing to suggest that £. formcae and E.
myrmecophaga are not distinct species.

ACKNOWLEDGEMENTS

I wish to express my deep gratitude ‘to R. P. Karf for
invaluable discussions about the nomenclatural problems of
validating these species, to D. Tyrrell and S. Batazy for

479

providing the type material of #. bullata and FE. formicae,
POs mmm GessmLOreconrecting the Latin descriptions, andsto
Heo epooper and DE. McCabe for their review and criticism
Dretheemanuscript.

LITERATURE CITED

BALAZY, S., and A. SOKOLOWSKI. 1977. Morphology and biology
of Entomophthora myrmecophaga. Trans. Britt. Mycol. Soc.
Bo melo 4-157,

DADOEIT, I:.J., C-W:. CROMPTON, and J.W. PARMALEE. 1978. An
atlas of atrborne pollen grains and common fungus spores
of Canada. Canad. Dept. Agric., Res. Branch (Ottawa),
Monograph No. 18.

BATKO, A. 1964. Some new combinations in the fungus family
Entomophthoraceae (Phycomycetes). Bull. Polon. Acad. Sct.
SCrmmcctUm btOlw ic: 405-406.

BATKO, A. 1966. On the subgenera of the fungus genus
Zoophthora Batko (Entomophthoraceae). Acta Mycol. 2:
15-21. :

BEN-ZE'EV, I., and R.G. KENNETH. 1981. Zoophthora ortenta-
lts sp. nov., a new fungal pathogen of Aphts cttricola
(Homoptera: Aphididae), and two new combinations of other
species of Entomophthoraceae. Phytoparastttca 9: 33-42.

BEN-ZE'EV, I., and R.G. KENNETH. 1982. Features-criteria of
taxonomic value in the Entomophthorales: I. A revision of
the Batkoan classification. In preparation for Mycotaxon.

FAEGRI, K., and J. IVERSEN. 1975. Textbook of pollen analy-
sts, 3rd ed. Hafner Press, New York.

HUMBER, R.A. 1975. Aspects of the biology of an insect-
parasitic fungus, Strongwellsea magna (Zygomycetes: Ento-
mophthorales). Ph.D. dissertation, University of Washing-
ton, Seattle.

HUMBER, R.A. 1976. The systematics of the genus Strong-
wellsea (Zygomycetes: Entomophthorales). Mycologia 68:
1042-1060.

HUMBER, R.A. 1981. An alternative view of certain taxonomic
criteria used in the Entomophthorales (Zygomycetes).
Mycotaxon 13: 191-240.

HUMBER, R.A. 1982. Strongwellsea vs. Erynta: the case for a
phylogenetic classification of the Entomophthorales (Zygo-
mycetes). In preparation for Mycotaxon.

HUMBER, R.A., and I. BEN-ZE'EV. 1981. Erynta (Zygomycetes:
Entomophthorales): emendation, synonymy, and transfers.
Mycotaxon 18: 506-516.

480

KRAMER, J.P. 1979. Interactions between blow flies (Calli-
phoridae) and Entomophthora bullata (Phycomycetes: Ento-
mophthorales). J. New York Entomol. Soe. 87; 135-140.

LOOS-FRANK, B., and G. ZIMMERMANN. 1976. Uber eine dem
Dicrocoeltum-Befall analoge Verhaltensanderung bei Amei-
sen der Gattung Formica durch einen Pilz der Gattung
Entomophthora. 2. Parasttenk. 49: 281-289.

MacLEOD sD eMan ep Deh RRELL eR Oe erOUP ER manda tem CCil len he
1973. Entomophthora bullata as a pathogen of Sarcophaga
alarichi. Jay TRVveErteDV mM IPaciGl i 2a<is/0-79-

POVAH, A.H.W. 1935. The fungi of Isle Royale, Lake Superior.
Papers Michtgan Acad. Set. Arts Let. 20: 113-156.

REMAUDIERE, G., and G.L. HENNEBERT. 1980. Révision systéma-
tique de Entomophthora aphtdts Hoffm. in Fres. Description
de deux nouveaux pathogénes d'aphides. Mycotaxon 11: 269-
Sle

REMAUDIERE, G., and S. KELLER. 1980. Reconsidération
systématique des genres d'Entomophthoraceae a potentialité
entomopathogéne. Mycotaxon 11: 323-338.

RIDDLE, L.W. 1906. On the cytology of the Entomophthoraceae.
Proc. Amer. Acad. Arts Sct. 42:

TURIAN, G., and J. WUEST. 1969. Mycoses a entomophthoracées
frappant des populations de fourmis et de drosophiles.
Mitt. Schwetz. Entomol. Ges. 42: 197-201.

TURIAN, G., and J. WUEST. 1977. Description complémentaire
de Zoophthora (Entomophthora) myrmecophaga Turian §& Wuest,
agent d'une mycose chez Servtformica fusca L. Mtt.
Sehwetz. Entomol. Ges. 50: 285-289.

YEN, D.F. 1962. An Entomophthora infection in the larva of
the tiger moth, Creatonotus gangis (Linnaeus). J. Insect
Pathol. 4: 88-94.

MYCOTAXON

Py One tee NOLO Dy 4 C5 05 October-December 1981

PRREVEEW SOE SLE
NONENTOMOGENOUS ENTOMOPHTHORALES

BeUuceet. B LUCKEY

Departinent of Botany, AJ-10
University of Washington
Seattle, WA 98195

SUMMARY.

The characters used to define genera in the nonentomogenous Ento-
mophthorales are evaluated and compared with those used for delimiting
entomogenous genera. It is proposed that the host organism should be
deemphasized. Five characters considered valid at the generic level
are 1) spore karyology; 2) sporophore architecture; 3) method of spore
discharge; 4) Zzygospore morphology and development; and 5) thallus
development. These characters are applied to the fifteen nonentomo-
genous fungi reported to date, and two taxonomic realignments are pro-
posed: Meristacrum pendulatum is transferred to Zygnemomyces and Ento-
mophthora vermicola is transferred to Macrobiotophthora. The transfer
of Tabanomyces milkoi to Meristacrum is accepted. Ancylistes netrii,
originally described without a Latin diagnosis, is validated. The ac-
cepted genera are: Ancylistes (3 sp-.), Ballocephala (3 sp.), Comple-
toria (1 sp.), Macrobiotophthora (2 sp.), Meristacrum (2 sp.), and Zyg-
nemomyces (2 sp.-). A key to the species of these genera is provided.

Doel ERODUCELON..

Entomophthorales are separated from other orders of
Zygomycotina by the forcible discharge of single spores and
a tendency to parasitize insects. There are, “nowever, sa
few saprobic species in Conidiobolus Brefeld and Basidio-
bolus MEIdaAmM An DIMNSa a. stalls sjroup .Of | species. Daran iti aug
noninsect' hosts. Humber, (1931s pp. 234)" observess atnat
Dewi neite 1Sseinappropriate. to, argue .abouGastiGe ChOlcemor
Cuitentamusedato. detine genera or -sboutmthewcorrectnescmor
one or another classification of the Entomophthorales with-
out taking the non-entomoyenous yenera into full account."
This paper attempts to render this accounting by reviewing
published records, type and liviny material of these fungi,
discussing the taxonomic criteria used to delimit the

482

1872.

1874.

1896.

1912.

1940.

1949.

19516

1952.

1968.

1970.

1973.6

1973.

1977.

O77.

1978.

1980.

TABLE 1.
REPORTS AND DESCRIPTIONS OF THE
NONENTOMOGENOUS ENTOMOPHTHORALES.

Ancylistes closterii Pfitzer, Monatsb. kKénigl. Preuss. Akad.

Wisse Berle 396.

Completoria complens Lohde, Tagebl. Versamml. Deutsch. Naturf.

Aertze 47: 206.

Ancylistes pfeifferi Beck, Verh. Zool. Bot. Ges. Wien 46: 233.

Macrobiotophthora vimariensis Reukauf, Centralbl. Bakt. Para-

site Infeckt. 63: 390.

Meristacrum asterospermum Drechsler, J. Wash. Acad. Sci. 30:

251.

*Ancylistes netrii Couch, J. Elisha Mitchell Sci. Soc. 65: 132.

Ballocephala sphaerospora Drechsler, Bull. Torrey Bot. Club 78:

199.

*Botryobolus parasiticus Arnaud, Bull. Soc. Mycol. Fr. 58: 184.

[=Ballocephala sphaerospora. ]

"A Conidiobolus-like fungus." Castaner, De Mycologia 60:
440-443.

tBallocephala verrucospora Richardson, Trans. Brit. Mycol. Soc.

55: 308.

Coelomomyces milkoi Dudka & Koval, Novitates System Plantarum

non Vascularium 10: 89.

Zygnemomyces echinulatus Miura, Rep. Tottori Mycol. Inst.

(Japan) 10: 520.

tEntomophthora vermicola McCulloch, Trans. Brit. Mycol. Soc. 68:

173.

+Meristacrum pendulatum McCulloch, Trans. Brit. Mycol. Soc. 68:

7 56

Ballocephala pedicellata Pohlad & Bernard, Mycologia 70: 131.

"Conidiobolus sp." Amin, E. & J. Webster, Trans. Brit. Mycol.
SOC 4 moo.

* Invalidly published due to lack of Latin diagnosis.
+ Type examined.

483

nonentomogenous genera, and proposing taxonomic revisions
where necessary.

There have been 14 entomophthoralean fungi reported
Since § 10/2 sfromesnoninsect ghosts, primarily ‘from “small
animals (nematodes, tardigrades) or plants in moist envi-
ronments (fern gametophytes, desmid algae). These fungi,
as well as the entomogenous Tabanomyces milkoi Couch et al.
f=Coetomomycesmmilkoil ~aresslisted pin Tables] in order of
Encl re publication. The useful data obtained from these
published descriptions, illustrations and photographs, and
from the original material are summarized in Tables 2-5.
Three nematophagous fungi are excluded from further consid-
eration in this paper. Two parasites of Heterodera Schmidt
(cyst-nematodes) were recently transferred to the Lagenid-
iales: Catenaria auxiliaris (Kuhn) Tribe ([=Tarichium
auxiliare Kuhn] MacLeod and Mlller-Kdgler 1970, Tribe 1980)
and Nematophthora gymnophila Kerry & Crump ([='Entomoph-
thora'-like fungus] Kerry 1974, Kerry and Crump _1980).
Haptoglossa Drechsler was placed in the Entomophthorales by
Ainsworth (1971), although Drechsler believed it belonged
in the Saprolegniales, a disposition supported by Davidson
ance baLeOnwmulg7 sa) wand Barronm (s90i0..

II. EVALUATION OF TAXONOMIC CHARACTERS.
Wee HOST.

Host range is often valuable as a taxonomic character
for delimiting nonentomogenous species since none have yet
been found from more than one host. In addition, nonento-
mogenous genera such as Ancylistes Pfitzer, Ballocephala
Drechsler and Completoria Lohde are restricted to one host
(see Table 2). However, as will be seen with the species
PlaceGamatmmvarlOusmecimesse: ingsgMeristacrumes Drechslenmea(M.
asterospermum, M. pendulatum, Tabanomyces milkoi), it is
poss iblejstor, natural relationships to be obscured if host
Characteristics are not tempered with an understanding of
basic aspects of morphology and development. The following
examination of these characterisitics provide a better
generic framework for the nonentomogenous Entomophthorales
than can be obtained from host characteristics alone.

2. THALLUS DEVELOPMENT.

The initial stages of infection are similar in most
nonentomogenous Entomophthorales, and involve the penetra-
f10n) Ob the cell wall ‘or. integument of the host by say thin
penetration tube arising from a spore (especially a second-
Buy spore) oreappressorium formed at the tip Of =a sterile

external hypha. Following penetration, the cytoplasmic
contents of the infective propayule migrates into a swollen
infection bladder. The thallus then ‘matures from®= this

bladder in one of three distinct patterns. In) the nematode
parasites (see Table 2), the thallus consists of long asep-
tate hyphae which more or less fill the host oryanism.
Maese hyphae do not form short cells or fragment into

484

TABLE 2.
HOST AND THALLUS CHARACTERISTICS IN THE
NONENTOMOGENOUS ENTOMOPHTHORALES.

INITIAL THALLUS MATURE THALLUS

SPECIES HOST form length x width form length x width
Ancylistes

closterii D AH FH x 7-5—11 SH 12.5-55 x 11-15

pfeifferi D AH FH x 7.5-14 SH 43-67 x 9-5-19

netrii D AH FH x 2-2-4.8 SH 15-30 x 8-15.5
Completoria

complens D HC 15=30* 7-15 same same
Macrobiotophthora

vimariensis TD (3 ? ? HB G ?

vermicola N AH FH 5-7 SH same
Castaner (1968) N AH FH 3-6-5 SH? same
Amin & Web.(1980) N AH FH @ SH? 18-23 x 12-18
Meristacrum

asterospermumt N AH 50-500 x 5-12 HB 25-6 05x 2

asterospermum® N AH

milkoi TB AH >mtz0e x 5-7 HB 70-176x 28-56
Zygnemomyces F

echinulatus N AH FH 2-5 same same

pendulatus N AH FH 5-12 same same
Ballocephala

sphaerospora TD AH 5-80 x 5=10 HB 40-80 x 5-10

verrucospora TD AH ; ie HB 20-40 x 10-12

pedicellata TD AH ? 2 HB 20-31 x 13-16
All measurements in pm. D=desmid algae AH=aseptate hyphae
° Davidson and Barron (1973) F=fern gametophyte HB=hyphal body
+ Drechsler (1940) N=nematode HC=hyphal cluster
* measured from illustration TB=tabanid larvae FH=filling host

TD=tardigrade SH=septate hyphae

hyphal bodies at maturity.
In Ancylistes, Ballocephala and Macrobiotophthora

vimariensis, after an initial mycelial stage, the hyphae

become regularly septate. The newly formed-cells may
remain attached, or fragment into freely floating hyphal
bodies, depending upon the physical activity of the host.

In Completoria, the infection bladder germinates and
forms numerous short hyphal lobes which form a radiating
cluster of cellsstiilVingsthe host cellsicompletely .alhiseis
reminiscent of growth patterns in some of the entomogenous
Entomophthorales (Thaxter 1888).

485

TABLE 3.
PRIMARY SPORE CHARACTERISTICS OF THE
NONENTOMOGENOUS ENTOMOPHTHORALES.

PRIMARY SPORE

SPECIES shape ornam length width base disch #spore devl
Ancylistes

closterii PYR - 13.6 5-17.25 P + 1 4h

pfeifferi PYR - 21-23.5 P + ay

netrii PYR - 16-25 P + 1 Be
Completoria

complens PYR = foe p == 1 T
Macrobiotophthora

vimariensis OBV - 18-21. 5x10-11.5* TR Fy 1 db

vermicola OBV = 16—2 26xXG9= 4 TReee 1 T
Castaner (1968) OV/B - 6.5-9.5x6.5-8.-5 TR + 1 ae
Amin & Web. (1980) OBV - 21-26 x 12-16 TR + i dW
Meristacrum

asterospermumt OBV = S= 119 xso-8 TR oh 5-15 B

asterospermum? OBV a I=17) - 5xO—10 TR ae 2-8 B

milkoi PYR - 12-16 x10-13 P + 4(3-1) B
Zygnemomyces

echinulatus SPH ECH 527-72 U = 10-20 A

pendulatus SPH ECH Da U = t=3 A
Ballocephala

sphaerospora SPH - 3-0-4.5 U + 20-80 A

verrucospora SPH ECH 4.5-6.0 U ae 4-12 A

pedicellata SPH - 6-7-8.0 U ate 2-10 A
All measurements in pm. ECH=Echinulate A=Acropetal
*Measured from published OBV=Obovoid B=Basipetal

illustrations. OV/B=Ovate-Obovate P=Papillate
° Davidson and Barron (1973) PYR=Pyriform T=Terminal
t Drechsler (1941) SPH=Spherical TR=Truncate

U=Unmodified

3. PRIMARY SPORES.
Morphology.

Shape. Three basic spore shapes are found in the non-
entomogenous Entomophthorales; all spores are of the uni-
tunicate type (Remaudiére and Hennebert 1980, Remaudiere
and Keller 1980). Ancylistes, Tabanomyces, and Completoria
have pyriform spores (i.e. globose spores with conical
Dapillace(berdaans.L939,.) Couch, 19497, Leitgeb s16o1)* ancmare
similar to those found in Conidiobolus. Meristacrum has
obovate spores with truncate bases (Davidson and Barron
LO Eee Anos tinal. y,s callocephala (Drechsler 1951, Rich-
ardson 1970, Pohlad and Bernard 1978), Zygnemomyces echin-
ulatus (Miura 1973) and Meristacrum pendulatum (McCulloch
1977) have globose spores lacking basal modifications.

486

Ornamentation. Spore ornamentation is uncommon in the
Entomophthorales. Some species of Massospora Peck emend.
Soper have verrucose spores, and the primary spores of Con-
idiobolus coronatus (Cost.) Batko may become villose by the
growth of appendages through the spore wall. Spore orna-
mentations consisting of small echinulations are also found
in Ballocephala verrucospora, Zygnemomyces echinulatus, and
Meristacrum pendulatum.

Karyology.

The morphological characterisitics of nuclei are con-
sidered important by Humber (1981). The presence or ab-
sence of nucleoli and heterochromatin (and hence the stain-
ing ability of the nuclei) are taxonomically useful at the
familial level (see §I1V.3), and certain genera with similar
spores (e.g. Conidiobolus, Entomophaga Batko) can best be
distinguished using nuclear morphology. The number of
nuclei per spore has been used to reveal the misplacement
of individual species (e.g. Culicicola Nieuwland [Humber
V98 15) yes

Unfortunately, few descriptions include karyological
data, although there are a few exceptions among the fungi
considered here. Hyphal bodies, gametangia and zygospores
of Ancylistes closterili apparently have several nuclei with
central nucleoli’ (Dangeard 1906). Couch (1949) was able’ to
Stain nuclei (weakly?) of A. netrii with lactophenol-cotton

blue and confirmed Dangeard's earlier reports. Moorman
(1976) also reported nucleoli in her study of mitosis in an
unidentified species of Ancylistes. From these limited

reports). it appears,) that sAncylistes Pis# mostaisimielanrm vo
Conidiobolus in terms of karyology (Humber pers. comm.)

Two nonentomogenous fungi, Completoria complens (Humber
pers. comm.) and Entomophthora vermicola, possess. the
heterochromatic nuclei lacking central nucleoli typical of
Entomophthora Fresenius.

4. PRIMARY SPOROPHORES.
Spore Number.

Most entomogenous species produce only one spore per
Sporophore. Species of the entomogenous genus Erynia Nowa-
kowski, while having apically branched sporophores, still
produce only one terminal spore per branch. The nonentomo-
genous members, however, can be divided into two groups.
Single-spored sporophores are found in Ancylistes, Complet-
Ooria, and Macrobiotophthora Reukauf. Pleuri- or multispor-
ed sporophores are found in Meristacrum, Tabanomyces Couch
et al., Zygnemomyces Miura and Ballocephala.

The number of spores in the multispored genera is
actually quite variable (see Table 3). Ballocephala has
two to 80 spores per sporophore, with most of this range
attributable to variation between individual sporophores of
B. sphaerospora. Similarly, Meristacrum asterospermum
usually produces five to 15 spores (Drechsler 1940) but was
found to produce as few as two (Davidson and Barron 1973).

487

Sporophore Septation and Spore Development.

Ballocephala produces sporophores with crosswalls at
the base of lateral sporiferous branchlets. The main axis
of the sporophore remains nonseptate, the spores are formed
terminally, and then are pushed aside as the sporophore
continues to elongate. Thus spores are formed and mature
in acropetal succession. Acropetal spore formation is also
Observed in Zygnemomyces echinulatus and Meristacrum
pendulatum (Figs. 1-2).

In Meristacrum asterospermum and Tabanomyces. milkoi
the sporophores become transversly septate, delimiting sep-
arate cells, each of which produces a single spore. Septum
formation, and hence spore maturation, is basipetal.

5. PRIMARY SPORE DISCHARGE MECHANISM.

Ingold (1934, 1971) distinguished three types of spore
discharge in the Entomophthorales: papilla-eversion (Conid-
iobolus), sporophore cannon (Entomophthora), and sporophore
rocket (Basidiobolus). The method of spore discharge found
in Ballocephala (see below) should be recognized as a
fourth type, and fungi that have lost spore discharge
could be placed in a fifth category.

The basic, and putative ancestral method of spore dis-
charge (papilla-eversion) involves the separation of the
wall layers of a columella which projects into the spore.
The domeshaped columella that remains (more or less intact)
after spore discharge and the everted conical or rounded
papilla at the spore base are diagnostic for this release
mechanism. Ancylistes, Completoria, and Entomophthora
vermicola (with simple sporophores), Tabanomyces milkol and
Meristacrum asterospermum (with septate sporophores) have
this type of spore discharge. The method of spore
discharge was not seen in the "Conidiobolus-like fungus"
(Castaner 1968) or Macrobiotophthora vimariensis, but can
be reasonably inferred from their spore shape. All other
spore discharge mechanisms are modifications of this basic
method.

The second type involves a sporophore having a more or
less swollen apex, and which ruptures just below the spore.
In Entomophthora s. str., the spore is campanulate, has a
flat base, and is carried away in a stream of sap shot from
the collapsing sporophore. It should be noted that in
Entomophthora muscae (Cohn) Fresenius, the secondary spores
are discharged by the papilla eversion method.

The third discharge mechanism, found in Basidiobolus,
involves the rupture of the base of a subspore swelling.
This swelling tapers apically to a point forming a minute
columella within the spore. The spore rockets away as the
swelling expels fluid from its torn end, and lands with or
without the remnants of the vesicle still attached to the
everted spore base. Theseclaimeathate tht Semismeid si tWwO-stage
rocket is probably more fanciful than accurate, as the
decrease in turgor pressure within the sporophore must be
accompanied by the simultaneous release of spore turgor by
the eversion of the papilla.

488

TABLE 4.
SPOROPHORE AND SECONDARY SPORE CHARACTERISTICS IN THE
NONENTOMOGENOUS ENTOMOPHTHORALES-

SPOROPHORE SUBSPORE CELL 2° SPORES

SPECIES length width} septn length width replic disch
Ancylistes

closterii 3-725 - - - ++ +

pfeifferi 4-7 = - - ae a

netrii 25=100°xX95S - - - + +
Completoria

complens 60-80 ? - - - - ?
Macrobiotophthora

vimariensis 4-5 - - - + -

vermicola 27-50 x 6-8 = - - + -
Castaner (1968) <60 X16. 5=3 - - - +* +
Amin & Web. (1980) <60-90 x ? - - - + +
Meristacrum

asterospermumt 200-500x6-9(3-5) +(S) - - + -

asterospermum® 60-500x5-9(3-5) +(S,U)- - + -

milkoi 90-200x7-8(4-6*) +(U) - - + -
Zygnemomyces

echinulatus 100-750x2.7-5.7 - 1-5-3 x «5-1-1 - -

pendulatus 18= 71x2-3 a 1=3 = =
Ballocephala

sphaerospora 50-175x4-5 = 4-7 x 3-4.5 = ~

verrucospora 50-150x5-7 - ew oe Mr - -

pedicellata 33— 9735 = 2-1 xX 1-6-2.6 = =

(1-1-1-6)

All measurements in pm. S=Spiralled sporophores
* measured from published illustrations U=Unspiralled sporophore
+ Drechsler (1941) ++=produces 2°+ 3° spores
° Davidson and Barron (1973) **=produces microspores

§ (narrowest width)

The sporophore discharge mechanism found in Ballo-
cephala constitutes a fourth type. All species of Ballo-
cephala have spherical spores lacking basal modifications,
but are subtended by a special cell (Separated from the
sporophore by a crosswall) whose rupture discharges the
spore. Drechsler (1951) proposed that this cell was, in
fact, the undischarged primary spore which produces a pre-
mature secondary spore, a hypothesis which is consistent
with the basic developmental patterns of the Entomophthor-
ales. In B. sphaerospora and B. verrucospora, the spore
bearing cells are utriculate, asymmetrical, curved upwards,
and as large as the spore (Drechsler 1941, Richardson
LOST MVD Vs Ballocephala pedicellata has ovoid to broadly
Obpyriform pedicels which are much smaller than the spores.
Spore discharge in this species is likely, although the
evidence is circumstantial (Pohlad and Bernard 1978).

489

Spore discharge is lacking in Zygnemomyces echinulatus
and Meristacrum pendulatum. In both species, the apex of
the sporophore gives rise to a terminal spore lacking basal
modifications, and then proliferates sympodially. The
resulting pedicels are not delimited by a crosswall,
although they may become devoid of cytoplasm. Thus, they
are of distinctly different origin, than the sporiferous
cells of Ballocephala.

6. SECONDARY" SPORES.

If forcible spore discharge is a key characterisitic

GfBcheseEnvcomophcthorales;—so too is the ability of thesdis—
charged spores to produce ‘secondary' spores (Table 4). In
almost every member of the order, primary spores that land
On hostile substrates can produce replicative spores which
aresetOorciblysdischarged. These spores are also called
repetitive, Or secondary (2°), tertiary (3%), etc. spores.
Exceptions to this rule are Ballocephala spp., Zygnemo-
myces echinculatus, and Meristacrum pendulatum.
Two additional types of ‘'secondary'" spores have been
taxonomically useful. Microspores ("multiplicative spores"
Drechsler 1952) are formed and discharged from single
primary spores of various species of Conidiobolus. Basid-
iobolus microsporus Benjamin produces elongate passively
detached "microspores" by a totally unrelated method (Ben-
jamin 1962, Tucker unpubl.). Microspores are also found in
the "Conidiobolus-like fungus" (Castaner 1968) but are
otherwise absent 1n nonentomogenous species.

Capillispores ("capilloconidia" Remaudiére and Henne-
bert 1980; "elongated secondary conidia" Drechsler 1955)
are usually slightly thick walled, ovoid, ellipsoid (or
Slightly irregular in shape), without modifications of the
spore base and borne on long slender sporophores ("capil-
lary conidiophores" Thaxter 1888). The loss of spore dis-
charge is apparently counterbalanced by the increased in-
fective potential or likelihood of dispersal by a passing
host. True capillispores are produced only by Basidio-
bolus. Nondischarged spores borne on tapering sporophores,
are found in Meristacrum asterospermum, Macrobiotophthora
vimariensis, and Entomophthora vermicola.

7. RESTING SPORES.
Morphology and Development.

Resting spores, whose function is one of perennation
rather than dispersal, are usually thick walled and formed
within the vegetative mycelium. The distinction between
zygospores (developing following conjugation) and azygo-
spores (developing without conjugation) has been questioned
(Humber 1981), but is here considered provisionally useful
(see below). Virtually all of the nonentomogenous Ento-
mophthorales produce resting spores of one form or another
(see Table 5).

Ancylistes, as in Conidiobolus, forms zygospores after
scalariform or lateral conjugation of unequal gametangia

490

TABLE 5.
RESTING SPORE CHARACTERISTICS OF THE
NONENTOMOGENOUS ENTOMOPHTHORALES

ZYGOSPORANGIUM ZYGOSPORE / AZYGOSPORE
SPECIES con form length width shape color orn diameter
Ancylistes
closterii U GB APPRESSED SPH BRN none 14. 5-20
pfeifferi U G 30-40 SPH BRN none 18. 5-22
netrii U cr. APPRESSED SPH fs none 19-26
Completoria
complens E? GB APP RESSED SPH - none 1 Wis
Macrobiotophthora
vimariensis - - - - - - -
vermicola E CE APPRESSED DDC - none 19-20
Castaner (1968) - - - - - -
Amin & Web.(1980) - - - - = - -
Meristacrum
asterospermumt - HB LOOSE SPH YEL UND 20
asterospermum® - HB LOOSE SPH UND T= 20
milkoi E G APP RESSED OVD ~ none 12-20x23-30
Zygnemomyces
echinulatus E HB/CT 22-28* OCT BRN none 16-24
pendulatus ? HB POL - none 18-24*
Ballocephala
sphaerospora E HB 28-33x14-16.5 DOL BRN none 17-20x13-16
verrucospora - - - - - - - - -
pedicellata E HB APPRESSED DOL BRN PUNC 20.6-26.6 x
17-3-18-6
All measurements in pm. con=conjugation (color)
*Measurements from pub=- E=equal gametangia BRN=Brown
lished illustrations. U=unequal gametangia YEL=Yellow
tDavidson & Barron (1973) dev=zygosporangia devel-
“Drechsler (1940) ope from or within: (shape)
CT=Conjugation tube DDC=Dodecahedral
orn=ornamentation G=Gametangium OVD=Ovoid
PUNC=punctate GB=Gametangial bud POL=Polygonal
UND=Undulate HB=Hyphal body SPH=Spherical

(PELtzer 81872))s. The presence of numerous protrusions on
the zygosporangial wall is used to separate A. pfeifferi

from other species of the genus (Berdan 1938, Couch 1949).

Completoria forms = one. toy.tens. (or. 20 )peroundmtom oval
thick walled resting spores within the enlarged cells in
the central portion of the hyphal mass. (The peripheral
cells of the thallus are by this time emptied by the forma-
tion of primary sporophores or infective hyphae.) Conjuga-
tion has not been observed (Atkinson 1884, 1885).

zygnemomyces echinulatus forms brownish, thick walled,
octangular zygospores following scalariform conjugation.
The octangular shape results from the position of the zygo-
Spore in the conjugation tube between the two parental

hyphal bodies (Fig. 1E). The resting spores of Meristacrum

491

pendulatumsy are, similarsin=shape (Fig. 2F), but the mode of
development is unknown.

Entomophthora vermicola forms zygospores after
scalariform or Lateral conjugation (Fig. 3K-P). At

maturity, these zygospores are dodecahedral, each face with
5 (4-6) raised edges and a central depression (Fig. 3P).

Conjugation in Ballocephala has not been proven,
although its occurrence was inferred by Drechsler (1951)
from the presence of two slightly inflated locules at
either end of the zygospore. The zygospore itself is
COMrormesOusecy Lindrical, sbrOwn atom dark—bDrown sine coLlor,
smooth or punctate, and is preferentially formed at reduced
temperatures (5° C) (Pohlad and Bernard 1978).

Meristacrum asterospermum was described by Drechsler
(1940) as forming azgyospores by the rounding up of the
cytoplasm of a thalloid segment, laying down of retaining
walls at either end, and deposition of a new undulate or
scalloped wall around the cytoplasm. The star-shaped rest-
ing spore is colorless or slightly yellowish, is about 15-
20 um in diameter, and contains a single large oil drop-
let.

Finally, in Tabanomyces milkoi, uninucleate, spherical
gametangia fuse in pairs with the contents of one gametan-
gium migrating into the other and forming a smooth, color-
less ovoid zygospore with a single diploid nucleus (Couch
et al. 1979, Nam and Dubitskii 1977).

zygospores vs. Azygospores: A Different Perspective.

Resting spores have been used for taxonomic purposes
to some extent, but recent treatments of the family
(Remaudiere and Keller 1980, Humber 1981) have cautioned
against some traditional applications of resting-spore
characteristics. In particular, the distinction between
zygospores and azygospores (with respect to conjugation)
was discussed at great length by Humber (1981), who. dismis-
sed it for two reasons. First, the development of (a)zygo-
Spores is often difficult to follow or interpret, and hence
published accounts may be misleading. This) Ob Jections is
valid, and the presence or absence of conjugation must be
used with caution. However, the second objection, that the
nuclear events of karyogamy and meiosis are of more import-
ance to the fungus than whether conjugation occurs, poses
an interesting question.

Of what importance are the sexual events of conjuga-
tion, karyogamy and meiosis to the entomophthoralean fungi?
It must first be noted that all "sexual" Entomophthorales
are homothallic (automictic) and homomictic (i.e. no mating
types exist). Conjugation, when it occurs, jis between
cells of the same branch or between hyphal bodies. Second-
ly, since most spores and hyphal bodies are multinucleate,
genetic recombination may occur somatically rather than
during meiosis. There are very few species in the Ento-
mophthorales for which evidence of meiosis exists (excep-
tions include Conidiobolus thromboides Drechsler= Entomoph-
thora virulenta Hall & Dunn, Tabanomyces milkoi). The lack
Of data on basic aspects of sexual events and genetics is a

492

great, hindrancel=ins attempts. lO, assess u Cicer iilpOLcaicemson
zygospores or to understand evolution within the order.

Retaining the ability to coevolve with host organisms
is jlikely; to be important sto =thesemtingl, but Ltesisrealso
apparent that the presence or absence of conjugation or of
karyogamy and meiosis do not seem to be as critical.
Perennation and infective ability, however, continue to be
important as a basic requirement of the entomophthoralean
Ti Lemcy Cle. The formation of the resting spore, whether
induced by conjugation or not, involves the phenotypic
expression of extensive portions of the genome, and thus
the shape, size, or ornamentation of resting spores may
ultimately be of more taxonomic significance than nuclear
SVCtl tors

8. SUMMARY OF TAXONOMIC CHARACTERS.

From the preceding discussion of morphological charac-
ters in the nonentomogenous Entomophthorales, it should be
clear that character diversity within the order is greater
than that encompassed by the entomogenous members alone.
Nevertheless, the three characters which Humber (1981) pro-
posed as most valuable in delimiting genera: "nuclear num-
ber wand (nature, " “sporophore branching; =| andy =.methiodmor
spore discharge," apply as well to the nonentomogenous
genera, and the assessment of these characters needs only
minor revision.

1. Spore karyology, i.e. nuclear morphology and num-
ber per primary spore, remains important in distinguishing
natural groups (i.e. families) within the Entomophthorales
(see §I1V.3). This may be of practical value for separat-
ing morphologically similar genera (e.g. Conidiobolus from
Entomophaga; or Ancylistes from Completoria).

2. Spore discharge 1s virtually universal within the
order and thus is presumably an ancestral character. The
Spore discharge "theme" has five variations: papilla-
eversion (Entomophthoraceae, Ancylistaceae), sporophore-
cannon (Entomophthoraceae), Ballocephala-type (Ancylist-
aceae), and sporophore-rocket (Basidliobolaceae), plus the
loss of discharge (Entomophthoraceae [Massospora], Ancyl-
istaceae [Zygnemomyces]).

3. Sporophore architecture now includes the presence
or absence of sporiferous branchlets or transverse septa-
tions as well as branching of the sporophore. In addition,
the mode of spore formation (basipetal, acropetal, or term-
inal) which is linked to sporophore septation, can be used
to distinguish genera. It is presumed that the unispored
sporophore is ancestral, and that multispored sporophores
are derived.

4. Zygospore morphology, which Humber (1981) dis-
counted but actually used as a secondary characteristic to
delimit Triplosporium (Thaxter) Batko, is most appropriate-
ly used for delimitation of species or subgenera.

III. GENERIC TAXONOMY AND REVISIONS.

493

ANGYEISTEOMPriLezer,
Monatsb. KoOnigl. Preuss. Akad. Wiss. Berlin. 396. 1872.

myper) Species; Ancylistes closterii Pfitzer, Monatsb.
Konigl. Preuss. Akad. Wiss. Berlin. 396. 1872.

Other species:

Ancylistes pfeifferi Beck, Verh. Zool.-Bot. Ges. Wien 46:
PEvemieec9'o.

Ancylistes netrii Couch, J. Elisha Mitchell Sci. Soc. 30:
251% 1949. Validation of this species is here
accomplished with the publication of the following
Latin diagnosis:

Hyphae asumentes primum aseptatae 2.2-4.8 pm crassae, tum ramosae et
septatae, cellulis 15-30 X 8-15 pm; hyphae exitibus unica in cellula
una factae, 2-4 pm crassae, parietem hospitis juxta tumidae usque ad
4-9 pm emergentes et hyphas externas facientes, quae hospitem proxi-
mum obnoxium inficunt, aut si ex aqua emergunt sporophora 25-100 ym
faccunt; sporae primigenae 16-25 pm crassae papillis conicis; zygo-
sporae intra hospitem inter aliquot hyphas longitudinales seriales
formatae, ut videtur conjugatione cellulis duabus hyphis oppositis,
zygosporis intra tubum conjugentem, sphaericus vel subsphaericis
19-26 pm crassis. Hospe: Netrium (Desmidiaceae).

Ancylistes is in many ways, Similar to Conidiobolus,
especially in spore shape and karyology (see §IV.2, IV.3).
The species are divided on the basis of host (Saccoderm or
placoderm desmid algae), size of the primary spore, and
zygosporangial morphology. See Berdan) (1938) “and? Couch
(1949) for a treatment of this genus.

BALLOCEPHALA Drechsler,
SOUL - BWeperathyy sendy (ONhisy RI Aer lee die

Type species: Ballocephala sphaerospora Drechsler, Bull.
Weyatente iweyen (agi sy 7Ase AU ile jonk:

Other species:

Ballocephala verrucospora Richardson, Trans. Brit. Mycol.
SOCHIES 3 7mO0 Cian LO 7 Oe

Ballocephala pedicellata Pohlad & Bernard, Mycologia 70:
Tsuba Cals ey,

Tivcmqenus;, =stound sonly sin Stardigrades ; miss unique min
the manner in which the spores are formed and discharged.
As previously mentioned (see §II.4, II.5), the discharged
spore may. actually be a secondary spore developing neoten-
ously from the undischarged primary spore, which is present
in the form of a utriculate vesicle or a reduced pedicel.
The species are separated on the basis of spore ornamenta-
tion and the morphology of the sporiferous cell.

494

COMPLETORIA Lohde,
Tagebl. Versamml. Deutsch. Naturf. Aertze 47: 206. 1874.

Type species: Completoria complens Lohde, Tagebl. Versamml.
Deutsch. Naturf£. Aertze 47: 206. 1874.

This rare parasite of fern prothalli has been reported
only twice since the original description (Leitgeb 1881,
Atkinson 1884, 1885). The presence of large stainable
nuclei lacking central nucleoli and a thallus composed of
short hyphal branches indicate a distant relationship with
most other nonentomogenous Entomophthorales. It is placed
in the Entomophthoraceae (see §IV.3).

MERISTACRUM Drechsler emend. Tucker & Humber,
Us Wash. (ACaAd oC) 30m 25 eer O40.

Type species: Meristacrum asterospermum Drechsler, J. Wash.
Negeiely Sebo ys, PGbhs AO.

Vegetative hyphae developing within living animals,
Simple to somewhat branched, at first continuous, later
dividing into disjointed segments which put forth a
sporophore into the air, the terminal portion dividing
basipetally, forming several to few cells, each of which
produces a Single spore forcibly discharged by papilla-
eversion; secondary spores replicative or passively detach-
ed; resting spores azygospores formed from hyphal segments
or zygospores formed from conjugation of spherical uni-
nucleate gametangia.

Other species:

Meristacrum milkoi (Dudka & Koval) Humber, Mycotaxon 13:

213 239 Galt.
Basionym: Coelomomyces milkoi Dudka & Koval apud Dudka,
cOval & Andreeva, Novitates Systematicae Plantarum

nonaVascularumelO<956 9-597 37.

Synonym: Pseudocoelomomyces milkoi Nam and Dubitskii,
Insts 2Z00l-mAlmawrAtayimAcadsii6cl). skazakhr woo Re Gv NEL
no. 1309-77 Dep.). 1977. (Lacking Latin diagnosis’)

Synonym: Tabanomyces milkoi (Dudka & Koval) Couch,
Andreeva’, / Laird’ Wx, \Nolani,;,. "Proc, Natl SAcad-mocis.
(U<S*A.:) 763" 230071979 SS i(Dericlentwebas tonympctitre)

Tabanomyces milkoi was transferred to Meristacrum by
Humber (1981) because of the Similarity in their sporophore
development. In M. asterospermum and M. milkoi,transverse
septa are layed down in basipetal succession, with each of
the newly formed cells producing a spore which is dis-
charged by papilla-eversion. This mode of sporophore
development is not found elsewhere in the Entomophthorales,
and is the single most important character in delimiting
the genus.

The difference in host organism is of questionable
value at the generic level, and the non-helicoid sporophore

495

of M. milkoi is of no significance since Davidson and Bar-
rons (19/3) "observed a simllar lack of spiralling) in sporo-—
phores emerging from resting spores of M. asterospermum.

There are, however, three differences between these
two species which may be used to divide Meristacrum into
subgenera when a fuller understanding of these and other as
yet undiscovered species is obtained.

Character M. asterospermum M. milkoi

1) Primary spore obovoid, globose
truncate base basal papilla

2) Secondary spore obovoid globose
passively detached replicative

3) Resting spores azygospores zygospores
spherical, undulate ovoid, smooth
conjugation (-) conjugation (+)

The resting spore characteristics are the most significant.

ZYGNEMOMYCES Miura emend.,
RED mELOUCOLIMMY CO) mins tamu apanmelO <a 2 0. med O10)7

Type species: Zygnemomyces echinulatus Miura,
ReEDemeLOLCOLIMMYCOl mi nctem(uapan)t 02520 Lod

Vegetative hyphae developing within living animal, un-
segmented, giving rise to sporophores; sporophores nonsep-
tate, bearing spores on pedicels formed from the sporophore
apex, proliferating sympodially at the apex or subapically;
primary spores globose, echinulate, produced acropetally,
not forcibly discharged; secondary spores absent; zygo-
spores thick walled, forming after lateral or scalariform
conjugation.

Other species:

ZYGNEMOMYCES PENDULATUS (McCulloch) comb. nov.,
Basionym: Meristacrum pendulatum McCulloch, Trans.
Bint femeMy.COumECOC MOG: ali > mero) Jie

Body of the infested nematode containing unsegmented
allantoid thalli, 2-5 pm wide; sporophcres erect, unseg-
mented, 18-71 pm long, after penetrating the host cuticle
forming an apical hook upon which a single terminal spore
develops, growth continuing by sympodial proliferation;
sterigmata formed from the sporophore hook, becoming
emptied of cytoplasm, 2-3.5 pm wide proximally, tapering to
0.8-1.2 pm wide at the spore; spore spherical, echinulate,
5-8 pm in diameter, adhering to the nematode when detached;
resting spores 18-24 pm, thick walled, with a single large
oil drop, found within dead nematodes.

496

ZYGNEMOMYCES ECHINULATUS Miura.

Fig. 1.

(A-E) Stages in sporophore development, showing acropetal maturation
of the spores; (F) zygospores formed following scalariform conjuga-
2E represents a portion of

tion. Redrawn from Miura
the iconotypus. Bar=10pm.

(1973). Fig.

497

ZYGNEMOMYCES PENDULATUS (McCulloch) Tucker.

Fige 2.

(A-E) Stages in sporophore development, showing the apical hook that
becomes the pedicel, and continued growth by sympodial proliferation;
(D-E) mature echinulate spores, the pedicel making an acute angle
with the sporophore base; (F) zygospores within the nematode. Drawn

from type slide. Bar=10pm.

498

Zygnemomyces was established by Miura (1973) who
distinguished Ze echinulatus from Ballocephala and
Meristacrum by three characters: echinulate spores (though
this is no longer valid with the discovery of Ballocephala

verrucospora by Richards [1970]), spores borne on pedicels

(and hence not discharged), and octangular zygospores

formed by scalariform conjugation. The first two
characters also are found in ae pendulatus, and a careful
reexamination of spore development has shown that both

species have sympodially proliferating sporophores (Figs. 1
& 2). The two species can be distinguished on the basis of
sporophore size (Table 4), spore number (Table 3), and
orientation of the pedicel.

The three multispored genera: Meristacrum, Balloceph-
ala, and Zygnemomyces, as here treated are coherent genera,
and each is quite easily distinguished from the others.
Meristacrum, with basipetally septate sporophores contrasts
with the nonseptate sporophores of Ballocephala and Zyg-
nemomyces. While spores develop acropetally in both of
these genera, the manner in which the spores are formed and
discharged (i.e., borne on propulsive sporiferous cells vs.
pedicels formed from the sporophore apex) is significantly
different.

MACRO BIOTOPHTHORA Reukauf emend.,
Centralbl.. Bak tewParasitt mintekt 2 Abe. 0 ly 6S roo Ue oie

Type species: Macrobiotophthora vimariensis Reukauf. Cen-
tralbl.) Bakt @ Parasit) SIntekt Ww AD tee W637 90 ee Ol.

Vegetative thallus developing internally within host,
giving rise to external hyphae and sporophores on which
Single terminal spores are borne; spores obovate, flat
based, forming secondary spores or germinating vegetative-
ly; conjugation scalariform or lateral forming zygospores,
resting spores absent, or producing chlamydospores.

This |genus was established by (ReukaulegCl 9 PZ) ee fOu uM.
vimariensis, a fungus which he found parasitizing Hypsibius

dujardini (Doyere) (=Macrobiotis lacustris Duj.) living

near a spring in Belvederer Parks in Weimar, Germany. This
fungus produced hyphal bodies which digested the fat bodies
of the tardigrade and then gave rise to a multitude of
sterile aseptate hyphae which penetrated the cuticle.
Primary spores were obovoid, uninucleate and were produced
terminally. These spores either germinated directly or
gave rise to secondary spores on slender’ sporophores.
While no other fungi similar to M. vimariensis have been
isolated from tardigrades, the following nematode parasite
is sufficiently similar to warrant transfer to this genus.

Other species:

499

MACROBIOTOPHTHORA VERMICOLA (McCulloch), comb. nov.,
Basionym: Entomophthora vermicola McCulloch. Trans.
Digiueme My .COlWmOOC O08 al/.3 ame OP TGS.0l—2 )i.ue lO? ye

Body of infested nematode filled with nonseptate
hyphae, 5-7 wm in diameter, developing erect hyphae which
penetrate the nematode cuticle; external hyphae hyaline,
unsegmented, flexuous (40-) 110-160 x 3-6 pm; sporophores
Py oURXmO-G NM, eStralght (Or bent at, the point of emergence
(OZ Us) aul oranched s(branchedtsaccordings to McCulloch):
primary spores terminal, obovoid, 16-22 x 9-14 pm (rarely
to 30 x 16 pm), pleurinucleate and unitunicate, discharged
at maturity, adhering to the cuticle of a passing nematode
and initiating new infections; secondary spores obovoid
medialy constricted, nonpapillate, 14.4-15.5 um long, 9-
iS soeaulleeineadiameterns narrowing at a ineck ™to= /-4—¢. 9° um;
secondary sporophores 17-20 pm long, 3.5-4.5 pm at base,
tapering to 1.8-2.2 wm at spore; zygospores 17-22 um,
dodecahedroid, faces centrally depressed with 4-6 edges.

Of the three type slides, only the one containing the
nematode illustrated in McCulloch (1977: Fig. 1) remains
DncaCt ma Ghigs., 3A~—J))% External hyphae are present, but
these are not the morphologically modified cystidia found
in species of Erynia (Remaudiére and Hennebert 1980, Humber
9S 1s) s Figure 3C shows the appearance of several of the
developing spores with a thin, hyaline outer wall layer
separated from the spore wall proper, and the: retraction of
the cytoplasm from the spore apex. While the separated
wall resembles the image seen in spores with bitunicate
walls, these effects are probably due to the lactophenol
mounting medium. The secondary spores are obovoid and
medially constricted, with a distinct neck below a more
globose region (Fig. 3H-J). A basal papilla is lacking
making secondary spore discharge unlikely. The sporophores
On which they are borne taper to a narrow point of attach-
ment, but they are not the true capillary sporophores of
Other genera.

Macrobiotophthora vermicola was originally placed in
Entomophthora, a genus to which it is no longer referable.
Living material of this species obtained from E. Bernard,
permitted additional characterization of the species, and
the following observations are in conflict with the origin-
aly description. Thies pramany SspOlese:contal na 4—O sin tomo—
phthora-type nuclei and are forcibly discharged by papilla
eversion. Secondary spores are medially constricted and
passively detached. Dodecahedral zygospores are formed
following scalariform or lateral conjugation.

Two Conidiobolus-like fungi reported from nematodes
(Castaner 1968, Amin and Webster 1980) are probably species
of Macrobiotophthora. However, nuclear morphology and num-
ber are unspecified, making placement of these two fungi
provisional. The method of thallus and primary spore form-
ation in all three nematophagous fungi is similar, but dif-
ferences include the absence of zygospores (although these
will probably be found), forcible discharge of secondary
spores, production of microspores (?), and variation in

500

SUL

spore Size (see Tables 2-5). Additional reports of para-
sites on terricolous animals may increase further the size
of Macrobiotophthora by at least two, if not more species.

IV. FAMILIES OF THE ENTOMOPHTHORALES.

At present, the order is variously divided, with one
family, Entomophthoraceae (Waterhouse 1973); two families,
Entomophthoracee and Basidiobolaceae (Alexopoulos and Mims
1979, Benjamin 1979); or three families, Entomophthoraceae,
Ancylistaceae, and Basidiobolaceae (Ubrizsy and V6érés 1966,
Batko 1974). Although three characters: karyology, spore
discharge mechanism, and conjuation might logically be used
to delimit these families, only karyology results in a
division consistent with current views of the order.

Lee Separation of the genera on the basis of spore
discharge method would result in four groups. The basic
papilla-eversion method has been modified in Entomophthora
to form a sporophore-cannon. The significance of this is
reduced, Since the secondary spores of E. muscae are still
dispersed by the papilla-eversion method. Samia ray,
Basidiobolus has a vestigial spore papilla, although the
propulsive force is provided by the rupture of a subspore
swelling.

Less obvious is the origin of the Ballocephala dis-
charge vesicle. The suggestion that the vesicle is homo-
logous with the primary spore (Drechsler 1951), and gives
rise directly to sessile secondary spores that (like capil-
lispores) are not dispersed, is consistent with the basic
developmental patterns of the Entomophthorales. Lt.also
should be noted that spore discharge was lost independently
by Massospora and Zygnemomyces. From the above observa-
tions, it can be concluded that spore discharge mechanism
cannot be used alone for family delimitation.

2ie Details of gametangial conjugation and resting
spore morphology are also possible criteria for family
delimitation. Zygospore characteristics have proven to be
useful and conservative within the other orders of the
Zygomycotina (Benjamin 1979). The Basidiobolaceae, with
its beaked gametangia, is distinct from all other members
of the order. The tendency of Conidiobolus and Ancylistes
to form zygospores in line with the unequal gametangla

Fige 3. MACROBIOTOPHTHORA VERMICOLA (McCulloch) Tucker.

(A-H) Spores and sporophores drawn from the type slide. (A-B) Devel-
opment of primary sporophores; (C) sporophore with retracted cyto-
plasm and separated outer wall layer; (D) primary spore with distinct
columella; (E) spore with everted papilla; (F) germinating primary
spore; (G-J) development of secondary spores, (I-J) medially con-
stricted obovoid secondary spores on tapering secondary sporophore.
(K-P) Conjugation and zygospores from living culture. Scalariform
(K) and lateral (L) conjugation; (M-N) developing zygospores; Mature
Zzygospore in optical section (0) and surface aspect (P).- Bar=10pm.

502

contrasts with the laterally budded zygospores of Entomo-
phthora. “Yet edivision, of the Order einto slaniliccmonmtiis
character alone is problematical due to the difficulties in
interpreting zygospore development in many species, and the
absence of conjugation in others. Until more research is
done on Entomophthorales, the utility of zygospore charac-—
ters will remain suspect.

oie Karyology is here regarded as the most useful
character, and three distinct lines can be delimited within
the Entomophthorales: Basidiobolus with large nuclei with
central nucleoli; Entomophthora, Entomophaga, Exrynila,
Strongwellsea Batko & Weiser emend. Humber, Massospora,
Completoria (Humber pers. comm.), and Macrobiotophthora
have small, readily staining nuclei with heterochromatin
but without a central nucleolus; and Ancylistes, Conidio-
bolus, Meristacrum, Ballocephala (?), and Zygnemomyces (?)
have small, poorly staining nuclei with a central nucleo-
lus. The elevation of these three groups to family status
is not only consistent with the currently accepted segrega-
tion of the Basidiobolaceae, but also unites those genera
which have similar spore discharge and zygospore character-
istics and provides a logical framework for the perception
of evolutionary trends within the Entomophthorales as a
whole.

V.- CONCLUDING REMARKS.

The taxonomic revisions proposed here are based upon
the evaluation of basic morphological characteristics.
Although important, the host organism is deemphasized at
the generic level. While research in the Entomophthorales
has been and probably will continue to be centered upon the
parasites of insects, the nonentomogenous fungi must not be
overlooked. It is hoped that as new’ reports of
nonentomogenous fungi are forthcoming, they will include
full descriptions of all aspects of nuclear morphology and
number, method of spore discharge, sporophore architecture,
zygospore development and morphology, and thallus
characteristics. Only then will a fuller understanding of
the order as a whole be obtained.

ACKNOWLEDGMENTS .

I would like to thank R. A Humber and R. K. Benjamin for reviewing
this article, and for their many comments, criticisms, and
encouragments; J. Ammirati for reading the manuscript; and D. Stuntz
for correcting the Latin diagnosis and proofreading the final draft.
Types were lent by the following institutions: Commonwealth
Mycological Institute (CMI) and Plant Pathology Herbarium, Dept. of
Primary Industries, Queensland (BRIP). E. Bernard kindly supplied the
new isolate of Macrobiotophthora vermicola.

503
i
KEY TO THE SPECIES OF NONENTOMOGENOUS ENTOMOPHTHORALES

we OPOLODNOres producing One. terminal Sporeeesecesseccacecececee
1". Sporophore producing several spores per sSporophoreseccecccce 7

2 e Spores IDV GASROL MN ohelelelersiorene’eleferere cictetoters cele ietere etererctereisis etetercietoretetttate
ON Spores ODOVat Ceres crovelercloic ec clceicie ele cierercicreteio crercioichete cvs erie ter teen 6

3 - Nuclei staining readily, parasitic on fern prothalli
eeocccccceccecccsvccccccccee ecOmpletoria complens
3". Nuclei poorly staining, parasitic on desmid algae...cccreeee 4

4. Zygosporangium with several protuberances; zygospore loose
within the zygosporangium........e.-eAncylistes pfeifferi
4'. Zygosporangium SMOG Mote orevere evellelel eo ololerere cle eiarcleherelererenereneresiorere ctene rs 5)

5 - Host Closterium (Placoderm Desmid); primary spores
13-17.5 PMecevvccccveccvcescccscccves eAncylistes closterii
5'. Host Netrium (Saccoderm); spores 16-25 pm....Ancylistes netrii

6 - Vegetative thallus consisting of hyphal bodies; parasitic on
tardigradesSecccccecceeseeeee eMacrobiotophthora vimariensis

6'. Vegetative thallus septate-mycelial; parasitic on nematodes
eeccccccceccvceseee eMacrobiotophthora vermicola

7 - Sporophores becoming transversly septate; spores developing
hbasipetallyeccccccccccccvvevcccccsccsccsesevesscesessseees 8

7'. Sporophores remaining nonseptate, or producing lateral,
basally septate sporiferous cells; spores developing
ACLOPeCALLYcccececvccccveccrscscccvcccevvcescesvssevessocs FY

8 . Spores pyriform; zygospores ovoid, punctate; 2° spores
YEPLicativeccecescccccecvvccccceccceee oMeristacrum milkoi

8'. Spores obovate; azygospores spherical, walls undulate; 2°
spores passively detached.......Meristacrum asterospermum

9 . Spores produced on basally septate sporiferous cells, the
rupture of which discharges the Sporeececccccscccevcee 10
9'. Spores produced terminally on the sporophore axiSececercesee 12

10 - Sporiferous cells utriculate, as large as spore or larger;
spores formed along entire sporophore axiSecerceccececee 11

10'. Sporiferous cells not inflated, smaller than spore; spores
clustered at the tip....eeeeeeeeeBallocephala pedicellata

11. Spores smooth, unornamented....+.e...+-eBallocephala sphaerospora
11'. Spores verrucoS€ecccccccsecevceceeseeeBallocephala verrucospora

12 . Sporophore apex straight, becoming transformed into a spore-
bearing pedicel forming an acute angle with the distal
end of the sporophore..-.eeeeeeeeeZygnemomyces echinulatus

12'. Sporophore apex becoming hooked, transformed into a pedicel
which forms an acute angle with the proximal end of the
SPOLOPhOLe oeececoescccesscssecesssuzygnemomyces pendulatus

504

LITERATURE CITED.

AINSWORTH, Ge C. 1971. Dictionary of the Fungi, 6th ed. Commonw.e
"Mycol. Inst., Kew. 663 pp.

ALEXOPOULOS, C. Je, and C. W. Mims. 1979. Introductory Mycology,
Third Eds John Wiley & Sons, New York. 632 pp.

AMIN, Ee. Ne Ee, and J. WEBSTER. 1980. British Records. Trans.
Brite Mycol. Soc. 74: 633-635.

ARNAUD, Ge 1952. Mycologie Concréte: Genera. Bull. Mycol. Soc. Fr.
58: 181-223.

ATKINSON, G. Fe. 1894. Completoria complens. Bot. Gaz. 19: 467-468.

ATKINSON, Ge Fe 1895. Damping Off. New York (State) Agr. Exp. Sta.
(Ithaca), Bull. 94: 231-272.

BARRON, Ge Le 1977. The Nematode Destroying Fungie Topics in Myco-
biol. noe 1. Canad. Biol. Publ.-, Guelph. 140 pp.

BATKO, A. 1974. Filogenza a struktury taksonomiczne Entomophthor-
aceaee In Ewolucja biologiczna: szkice teoretyczne i metodol-
ogiczne He, Nowinskiego, ed.-)- Polska Akad. Nauk, Inst. Fil-
ozof. Socjol., Wroclaw.

BECK, Ge 1896. Ancylistes Pfeifferi ne. sp.e Verhandl. Zool.-Bot.
Gese Wien. 233.

BENJAMIN, Re Ke 1962. A new Basidiobolus that forms microspores.
Aliso 5: 223-233.

BENJAMIN, Re Ke 1979+ Zygomycetes and their spores. pp 573-621 in
Kendrick, Be (ed-). The Whole Fungus: The sexual-asexual
synthesis. National Museums of Canada. Ottawa. 1979.

BERDAN, H. 1938. Revision of the Genus Ancylistes. Mycologia 30:
396-415.

CASTANER, De 1968. A Conidiobolus-like fungus destroying nematodes
in Iowa. Mycologia 60: 440-443. :

COUCH, J. N. 1949. A new species of Ancylistes on a saccoderm
desmid. J. Elisha Mitchell Sci. Soc. 65: 131-136.

COUCH, Je. Ne, Re Ve ANDREEVA, M. LAIRD, and Re A. NOLAN. 1979.
Tabanomyces milkoi (Dudka and Koval) emended, genus novum, a
fungal pathogen of horseflies. Proce Natl. Acad. Sci. U.S.A.
703° 2299=2302.

DANGEARD, Pe. Ae 1906- Recherches sur le développement du périthéce
chez les Ascomycetes. Le Botanist 9: 157-303.

DAVIDSON, Je Ge Ne, and Ge Le BARRON. 1973. Nematophagous fungi:
Meristacrum. Canad. J.» Bote 51: 231-233.

DAVIDSON, J» Ge Ne, and Ge Le BARRON. 1973a- Nematophagaous fungi:
Haptoglossa. Canad. J- Bot. 51: 1317-1323.

DRECHSLER, Ce 1940. Three fungi destructive to free-living terric-
Olous nematodes. J. Wash. Acad. Sci. 30: 240-254.

DRECHSLER, C. IRIE An entomophthoraceous tardigrade parasite
producing small conidia on propulsive cells in spicate heads.
Bull. Torrey Bot. Club 78: 183-200.

DRECHSLER, C. 1955. A small Conidiobolus with globose and with
elongated secondary conidia. J. Wash. Acad. Sci. 45:114-117.

HUMBER, R. A. 1981. An alternative view of certain taxonomic cri-
teria used in the Entomophthorales (Zygomycetes). Mycotaxon
13: 191-240.

INGOLD, C. T. 1934. The spore discharge mechanism of Basidiobolus
ranarume New Phytol. 33: 274-277.

905

INGOLD, C. Te. 1971. Fungal spores: their liberation and dispersal.
Claredon Press. Oxford.

KERRY, Be Re 1974. A fungus associated with young females of the
cereal cyst-nematode Heterodera avenae. Nematologica 20:
259-260.

KERRY, Be Re, and De He CRUMP. 1980. Two fungi parasitic on females
of cyst-nematodes (Heterodera spp.). Trans. Brit. Mycol. Soc.
74: 119-125.

LEITGEB, M. He. 1881. Completoria complens Lohde, ein in Farnpro-
thallien schmarotzender Pilz. Sitzungsber. Akad. Wiss.,
Math.-Naturwiss. Cle, Abt. 1. 84: 288-324.

LOHDE, G.- 1874. Uber einige neue parasitische Pilze. Tagebl. Ver-
samml. Deutsch. Naturf. Aerzte 47: 203-206.

MACLEOD, D. Me, and Es. MULLER-KOGLER. 1970. Insect pathogens:
species originally described from their resting spores mostly
as Tarichium species (Entomophthorales: Entomophthoraceae).
Mycologia 62: 33-66.

MCCULLOCH, Je Se 1977. New species of nematophagous fungi from
Queensland. Trans. Brite Mycol. Soc. 68: 173-179.

MIURA, Ke 1973. Three entomophthoralean parasites of nematodes
collected from Japan. Rep. Tottori Mycol. Inst. (Japan) 10:

517-522.
MOORMAN, Ge We 1976. Mitosis in Ancylistes spe Mycologia 68: 902-
909.

NAM, E. A., and A. M. DUBITSKII. 1977. A review of the systematic
status of the fungus Coelomomyces milkoi and a description of
a new genus of the entomophthorous fungus Pseudocoelomomyces
(Entomophthorales). Inst. Zool. Alma Ata Acad. Nauk Kazakh.
SSR. (VINITI no. 1309-77 DEP., UDK 591.69-577.1:576).

PFITZER, Ee 1872. Ein neuer Algen-Parasit aus der Ordnung der
phycomyceten. Monatsb. Konigl. Preuss. Akad. Wiss. Berlin.
379-398.

POHLAD, Be. Re, and E. C. BERNARD. 1978. A new species of Entomoph-
thorales parasitizing tardigrades. Mycologia 70: 130-139.

REMAUDIERE, Ge, and G. C. HENNEBERT. 1980. Révision systématique de
Entomophthora aphidis Hoffm. in Fres. Description de deux nou-
veaux pathogénes d'aphides. Mycotaxon 11: 269-321.

REMAUDIERE, Ge, and S. KELLER. 1980. Revision systématique des
genres d'Entomophthoraceae a potentialité entomopathogene.
Mycotaxon 11: 323-338.

REUKAUF, Ee 1912. Ein verderber des Wasserbadren Macrobiotus lacus-
tris Duj., Macrobiothophthora vimariensis (Reukauf). Central-
bl. Bakt. Parasite. Infekt. Abte 1, 63: 390-393.

RICHARDSON, Me. Je 1970. Ballocephala verrucospora Spe nove, para-
sitizing tardigrades. Trans. Brit. Mycol. Soc. 55: 307-309.

SOPER, Re Se 1974. The genus Massospora, entomopathogenic for cica-
das. Part Ie Taxonomy of the genus. Mycotaxon 1: 13-40.

THAXTER, Re 1888. The Entomophthoreae of the United States. Mem.
Boston Soc. Nature Hist. 4: 133-201.

TRIBE, He Te. 1977. A parasite of white cysts of Heterodera: Caten-
aria auxiliaris. Trans. Brit. Mycol. Soc. 69: 367-376.
UBRIZSY, Ge, and Je VOROS. 1966. A new conception in the review on
the phylogeny and system of fungi. Acta Botan. Acad. Sci.

Hunge 12: 199-220.

MYCOTAXON

Vols) XL Bie NOt opp D eS UO sono October-December 1981

ERYNIA (ZYGOMYCETES: ENTOMOPHTHORALES ) :

EMENDATION, SYNONYMY, AND TRANSFERS

RICHARD A. HUMBER

USDA-SEA-ARS Insect Pathology Research Untt
Boyce Thompson Instttute at Cornell
Tower Road, Ithaca, New York 14853, USA

DS RAG IG BN beers,

NSERC/Agrtculture Canada, Research Branch
Researeh Statton Harrow
Harrow, Ontarto NOR 1G0, Canada

SUMMARY

The emended sense of Erynia Nowakowski (Entomophthor-
ales: Entomophthoraceae) is limited to species having uni-
nucleate, bitunicate primary conidia borne on (digitately-
and apically) branched conidiophores and forcibly discharged
by the eversion of the basal papilla. This emendation spe-
cifically (1) rejects a separation of Zoophthora Batko from
Erynta based on the presence of capilliconidia or any other
criterion, (2) rejects any synonymy of Strongwellsea Batko
& Weiser emend. Humber with Zrynta, and (3) excludes from
Erynta any species having conidia which are multinucleate,
unitunicate, or borne on simple (or rarely or basally
branched) conidiophores. Zoophthora Batko is rejected as a
later synonym of Erynta Nowak., and the remaining species
of Zoophthora not previously placed in Erynta are trans-
ferred. All recognized species of Erynta sensu Humber &
Ben-Ze'ev are cited.

The recognition of two genera, Zoophthora Batko (1964a)
and Erynta Nowakowski (1881), separated principally by the
formation of capilliconidia (secondary conidia formed atop
Capillary conidiophores) by Remaudiére and Hennebert (1980)
has raised a problem in synonymies (Humber 198la): If the
type species of Zoophthora, Z. radtcans (Brefeld) Batko (=

507

Entomophthora sphaerosperma Fres. sensu Thaxter 1888), and
the type species of Erynta, EF. ovtspora (Nowakowski) Nowak-
Owski, are accepted on any basis to be in distinct genera,
then both of these generic names may stand. If, however,
one accepts Batko's sense of Zoophthora for all species with
uninucleate, bitunicate conidia borne on branched conidio-
pnores (Batko 1964a-b, 1966a-b, 1974; Humber 198la-b, 1982;
Ben-Ze'ev and Kenneth 198la-b, 1982a-b), then both of these
species must be in the same genus. Further, the Interna-
tional Code of Botanical Nomenclature (ICBN) requires that
Batko should have adopted the older generic name, Erynta,
rather than propose a new name for this large group of
Species.

Humber (198la-b; 1982), Ben-Ze'ev (1980), and Ben-Ze'ev
and Kenneth (198la-b, 1982a-b) find no basis for the sepa-
rate and simultaneous usage of Zoophthora and Erynta espoused
by Remaudiére and Hennebert (1980) and Remaudiére and Keller
(1980). Among the possible resolutions of this synonymy, the
only one that is both in accord with the ICBN and the prior
transfer of most Zoophthora species to Erynta (Remaudiére and
Hennebert 1980; Humber 198la-b) is now to take up £Zrynta
Nowakowski as the correct name of the genus which includes
Entomophthora radtcans Brefeld. Accordingly, we present an
emended circumscription for Erynta, and regard Zoophthora to
be a younger synonym.

ERYNIA Nowakowski emend. Humber & Ben-Ze'ev, emend.
nov.

Erynta Nowakowski, 1881, Datenn. III Zjazdu Lek.
Przgyr. Polak. Krakow., Seke. Bot. 6, 67-68.

Zoophthora Batko, 1964, BULL ACAd aeRO LON weoCt.
(NOs (Nei, Meh, MAL Gas.

VEGETATIVE STRUCTURES usually hyphal bodies or mycelium, but
exact form depends upon stage of development. NUCLEI easily
observed, 4.5-15 um diameter; without a prominent central
nucleolus; filled with conspicuous heterochromatic threads
or granules. CONIDIOPHORES apically branched, forming a
(usually digitate) cluster of conidiogenous cells; each
branch basally septate, each conidiogenous cell usually uni-
nucleate, producing a single apical conidium; some conidio-
phores unbranched, producing a single terminal conidiogenous
cell and one conidium. PRIMARY CONIDIA uninucleate, bituni-
cate (with a separable outer wall layer except over the basal
papilla); shape variable but usually ovoid, clavate, turbi-

508

nate, or elongate, radially symmetrical or curved; papilla
centered on spore axis or displaced laterally; forcibly dis-
charged by eversion of the papilla against the conidiophore.
SECONDARY CONIDIA more or less similar to primary conidia
(or with short radiate arms), formed singly on a short coni-
diophore arising from a primary conidium and forcibly dis-
charged by eversion of the papilla or else amygdaliform to
elongate, clavate, falcate, or lanceolate, usually curved,
and borne on an extended, thin capillary conidiophore aris-
ing from a primary or secondary conidium. RESTING SPORES
usually globose with two thickened wall layers; epispore
colorless or variously colored, smooth or variously orna-
mented; budded laterally or terminally from parental hyphal
body whether as zygospores or azygospores. RHIZOIDS present
in most species, either (1) hypha-like, simple or branched,
solitary or aggregated into pseudorhizomorphs, terminating
in more or less branched (but only rarely discoid) holdfast
of (2) stout, unbranched, solitary or loosely fasciculate
(but not aggregated in pseudorhizomorphs), terminating in a
richly branched discoid or irregularly expanded holdfast;
rhizoids may not form when resting spores are produced.
CYSTIDIA absent in most species; simple, usually aseptate,
Spear-like, extending slightly above level of conidiophores
or else thick and columnar, aseptate, usually branching api-
cally, extending far beyond level of conidiophores; cystidia
usually absent when resting spores are formed in tne host.

TYPE SPECIES: Erynta ovtspora (Nowakowski) Nowakowski,
1881, Datenn. IIT Zjazdu Lek. Prayr. Polak. Krakéu.,
Seke. Bot. 6, 67-68. Basionym: Entomophthora ovtspora
Nowakowski, 1877, Bot. Zettg. (Letpzig) 35, 20.

This emendation rejects Zoophthora as a separate genus
for species forming passively detached secondary capilli-
conidia as advocated by Remaudiére and Hennebert (1980).
It also rejects a definition of Hrynta based on conidial
morphology (Remaudiére and Hennebert 1980; Remaudiére and
Keller 1980), thereby excluding Erynta carolintana (Thax.)
Rem. §& Henneb. (with unitunicate, plurinucleate conidia on
simple conidiophores) and restoring Strongwellsea Batko §
Weiser emend. Humber (with bitunicate, uninucleate conidia
on unvaryingly simple conidiophores; cf. Remaudiére and
Keller 1980, Humber 198la, 1982). This emendation also
acknowledges the formation of multiradiate secondary coni-
dia by some riparian species (Descals et al. 1981).

509

The formal adoption of Erynta as the nomenclaturally
correct name for the genus circumscribed above requires the
transfer of all remaining names placed in Zoophthora but not
yet formally acknowledged as species of Frynta.

ERYNIA APHIDIS (Hoffmann in Fresenius) Humber § Ben-Ze'ev, comb. nov.
BASIONYM: Entomophthora aphidis Hoffmann in Fresenius, 1856, Ab-
handl. Senekenberg. naturf. Ges. 2: 208 emend. Remaudiére § Henne-
bert, 1980, Myecotaxon 11: 290, NON Entomophthora aphtdis Hoffm. in
Fres. sensu Thaxter, 1888, Mem. Boston Soc. Nat. Htst. 4: 175.

ERYNIA CANADENSIS (MacLeod, Tyrrell, § Soper) Humber § Ben-Ze'ev,
comb. nov. BASIONYM: Entomophthora canadensis MacLeod, Tyrrell, §&
Soper. 19/9, Candd..d. (Bot, 67: 2664.

ERYNIA CRASSITUNICATA (Keller) Humber § Ben-Ze'tev, comb. nov.
BASIONYM: Zoophthora crassttuntcata Keller, 1980, Sydowta Ann.
MYCOU. OCP a lan moos 1102

ERYNIA COLEOPTERORUM (Petch) Humber §& Ben-Ze'ev, comb. nov. BASIO-
NYM: Entomophthora coleopterorum Petch, 1932, Trans. Brit. Mycol.
OCm ue CiaRL es

ERYNIA CRUSTOSA (MacLeod §& Tyrrell) Humber §& Ben-Ze'ev, comb. nov.
BASIONYM: Entomophthora crustosa MacLeod §& Tyrrell, 1979, Canad.
Enconmol edd sel 138.

ERYNIA ELATERIDIPHAGA (Turian) Humber, Ben-Ze'ev, §& Kenneth, comb.
nov. BASIONYM: Entomophthora elateridtphaga Turian, 1978, Mtt.
Schwetz. Entomol. Ges. 51: 398.

ERYNIA GEOMETRALIS (Thaxter) Humber § Ben-Ze'ev, comb. nov. BASIO-
NYM: Empusa (Entomophthora) geometralts Thaxter, 1888, Mem. Boston
DOC NOL BOLSC. a4 ce 1/0.

ERYNIA HENRICI (Molliard) Humber § Ben-Ze'ev, comb. nov. BASIONYM:
Entomophthora henrict Molliard, 1918, Compt. Rend. Hebd. Séances
ACG OCL en earre 16757958;

ERYNIA JACZEWSKII (Zaprometov in Jaczewski § Jaczewski) Humber, comb.
nov. BASIONYM: Tartehtum jaczewskitt Zaprometov in Jaczewski § Jac-
zewski, 1931, Opredelttel grtbov 1: 225.

ERYNIA LANCEOLATA (Keller) Humber § Ben-Ze'ev, comb. nov. BASIONYM:
Zoophthora lanceolata Keller, 1980, Sydowta Ann. Mycol., Ser. II,
6059 155%

ERYNIA NEBRIAE (Raunkiaer) Humber & Ben-Ze'tev, comb. nov. BASIONYM:
Entomophthora nebriae Raunkiaer, 1892, Bot. Tidsskr. 18: 108-111.

ERYNIA OCCIDENTALIS (Thaxter) Humber §& Ben-Ze'ev, comb. nov. BASIO-
NYM: Empusa (Entomophthora) occidentalts Thaxter, 1888, Mem. Boston
DOC MMIC EU BHESt ee al7.1.

ERYNIA ORIENTALIS (Ben-Ze'ev & Kenneth) Humber, Ben-Ze'ev, & Kenneth,
comb. nov. BASIONYM: Zoophthora ortentalis Ben-Ze'ev §& Kenneth,
1981, Phytoparastttca 9: 35.

ERYNIA PHALLOIDES (Batko) Humber § Ben-Ze'ev, comb. nov. BASIONYM:
Zoophthora phalloides Batko, 1966, Acta Mycol. 2: 7.

ERYNIA PHYTONOMI (Arthur) Humber, Ben-Ze'ev, & Kenneth, comb. nov.
BASIONYM: Entomophthora phytonomt Arthur, 1886, Bot. Gaz. (Craw-
FOVUSeL Lie) s11si4.

ERYNIA RADICANS (Brefeld) Humber, Ben-Ze'ev, § Kenneth, comb. nov.
BASIONYM: Entomophthora radicans Brefeld, 1870, Bot. Zeitg.
(Letpaztg) 28: 186.

510

Entomophthora coleopterorum Petch (1932) is included in
the above list because of the conjunction of long, narrowly |
oval conidia (32-44 x 8-14 um), two fascicles of stout rhi-
zoids emerging from specific points on the ventral thorax,
and cystidia (Petch 1944). This set of characters occurring
together is known only from Hrynta species although it must
still be determined that £. coleopterorum has uninucleate,
bitunicate conidia and branched conidiophores. Petch (1944)
regarded this species to be a probable synonym of Entomoph-
thora carpenttert Giard (1888) whose description was based
wholly on the unique mode of rhizoidal attachment of affect-
ed elaterid beetles. Turian (1957) identified as &. car-
penttert a fungus he found on beetles which died head-down
and were attached to the plant by two fascicles of rhizoids;
he claimed this fungus produced uninucleate, globose conidia.
Humber (198la) raised doubts about Turian's interpretation
and discussed the possible synonymy of F£. carpenttert and E.
coleopterorum. Until any extant material of these collec-
tions can be re-examined, there is no doubt that the best
characterized of these collections is FE. coleopterorum, and
that this species is more certainly a species of Erynta than
of any other entomophthoralean genus.

Entomophthora henrtct Molliard (1918), which was de-
scribed from mosquitoes (Culex pipiens), 1S accepted here as
an Erynta species by virtue of its much branched conidio-
phores, presence of cystidia (which Molliard stated were also
produced in culture), and of rhizoids with discoid or irregu-
larly ramified holdfasts. It remains to be established
whether the conidia of this species are uninucleate and bi-
tunicate as may be predicted from this constellation of other
characters. Hrynta henrict resembles EF. ovtspora, but insuf-
ficient information about the morphological plasticity of
either species currently €xistsPto supports thespropost trons,
Waterhouse (1975) of their synonymy.

Entomophthora nebriae Raunkiaer (1892), which attacks
carabid beetles (Webria brevtcolis), has elongate conidia
(28.0-37.0 x 10.0-13.0 um) borne on richly branched conidio-
phores. Rhizoids are present, but their morphology is not
described. No secondary conidial forms or cystidia are men-
tioned by Raunkiaer (1892). The resting spores of this spe-
cies are formed outside of the host body, and are spherical,
36-50 wm in diameter, with a pale brown, smooth, and rather
thin wall. Batko (1966b) regarded this species to belong in
Zoophthora subg. Zoophthora.

Zoophthora jacezewskit (Zaprometov in Jaczewski §& Jaczew-
Ski) Batko (1964b) was not discussed by Remaudiére and col-

leagues in their revision of entomophthoralean taxonomy. We
have been unable to obtain the literature to confirm either
the nomenclatural validity of Tartchtum jaczewskit Zaprometov
in Jaczewski § Jaczewski (1931; see also MacLeod and Muller-
Kogler 1970) or the appropriateness of Batko's acceptance of
Entomophthora sabrit Rozsypal (1951; an invalidly published
nomen nudum) as the conidial state of T. jaczewskit. One of
the present authors (RAH) feels that it is not unreasonable
to accept both Batko's acknowledgement of 7. jaczewskit to
be validly published and of his synonymization of £. zabrit
with this species.

Those species previously described as or transferred to
Erynta and accepted according to the above emendation in-
clude the following:

Erynta amertcana (Thaxter) Remaudiére §& Hennebert (1980)

Erynta aquattca (Anderson & Ringo ex Anderson § Anagnosta-
kis) Humber (198la) non Zrynta aquattca (Anderson § Ringo)
Remaudiére § Hennebert (1980)

Erynta blunktt (Lakon ex Zimmermann) Remaudiére §& Hennebert
(1980)

Erynta brahminae (Bose & Mehta) Remaudiére §& Hennebert (1980)
(possibly synonymous with Lrynta echtnospora)

Erynta bullata Thaxter §& MacLeod in Humber (1981b) non Erynta
bullata (Thaxter in Povah) Remaudiére §& Hennebert (1980)

Erynta calltphorae (Giard) Remaudiére & Hennebert (1980)

Erynta eontca (Nowakowski) Remaudiére § Hennebert (1980)

Erynta ereatonott Yen in Humber (1981b) non E£rynta creato-
nott (Yen) Remaudiére §& Hennebert (1980)

Erynta curvtspora (Nowakowski) Nowakowski (1881)

Erynta delptntana (Cavara) Humber (1981a)

Erynta delphacts (Hori) Humber (1981la)

Erynta diptertgena (Thaxter) Remaudiére §& Hennebert (1980)

Erynta echinospora (Thaxter) Remaudiére §& Hennebert (1980)

Erynita erinacea (Ben-Ze'ev §& Kenneth) Remaudiére §& Hennebert
(1980)

Erynta formtcae Humber §& Batazy in Humber (1981b)

Erynta gloeospora (Vuillemin) Remaudiére §& Hennebert (1980)

Erynta gractlts (Thaxter) Remaudiére §& Hennebert (1980)

Erynta montana (Thaxter) Remaudiére & Hennebert (1980)

Erynta myrmecophaga Turian & Wuest in Humber (1981b) non
Erynta myrmecophaga (Turian §& Wuest) Remaudiére §& Henne-
bert (1980)

Erynta neoaphidts Remaudiére §& Hennebert (1980) [= Entomoph-
thora aphtdis Hoffmann in Fresenius sensu Thaxter (1888) ]

Erynta nouryi Remaudiére §& Hennebert (1980) [= Entomophthora

5i2

exitttalts Hall §& Dunn sensu Gustafsson (1965) ]
Erynta ovtspora (Nowakowski) Nowakowski (1881)
Erynia phalangtetda (Lagerheim) Remaudiére § Hennebert (1980)
Erynita rhtzospora (Thaxter) Remaudiére § Hennebert (1980)
Erynia sepulchralis (Thaxter) Remaudiére §& Hennebert (1980)
Erynta vartabtlts (Thaxter) Remaudiére § Hennebert (1980)
Erynta virescens (Thaxter) Remaudiére §& Hennebert (1980)
Erynta vomttortae (Rozsypal) Remaudiére & Hennebert (1980)

Doubtful or) excluded species.

Erynta carolintana (Thaxter) Remaudiére §& Hennebert (1980)
= Entomophthora carolintana (Thaxter) Keller, 1978, Sydow-
ta Ann. Mycol., Ser. II, 31: 88 = Empusa caroltntana Thax-
terl 188Syilen. BOSton pOc mNOe. HUScem love

Erynta castrans (Batko §& Weiser) Remaudiére & Keller (1980)
= Strongwellsea castrans Batko & Weiser, 1965, J. Invert.
Pathol. 14: 463 emend. Humber, 1976, Mycologia 68: 1056.

Erynta magna (Humber) Remaudiére & Keller (1980) = Strong-
wellsea magna Humber, 1976, Mycologta 68: 1057.

Zoophthora extttalis (Hall & Dunn) Batko, 1966, Bull. Polon.
AGadieSets, eoecryusct. Biola io 40 Sma pmeiLomopiT ora
exttralts Hall & Dunn, 1957, Htlgardta 27: 163,

Zoophthora ferruginea (Phillips in Houghton & Phillips)
Batko, 1966, Acta Mycol. 2: 19 = &ntomophthora ferrugtnea
Phillips in’ Houghton @ Phillips, 1886, Ari. Mao. Nacvweeer.
Ser. 5, 18: 6. |

Zoophthora forftculae (Giard) Batko, 1964, Bull. Potlon. Acad.
Sev. , Sen. SCt.Biol. “12: A040= | Empusasforficulae(Gianw
Petch, 1944) Trans. Broce Mycol. WSOC. 427-8 =e LOmO pi
thora forftculae Giard, 1889, Bull. Set. France Belgtque
AOS PANE

Empusa carolintana and the two species of Stronguellsea
are excluded from Zrynta for reasons mentioned above and
discussed extensively by Humber (198la, 1982).

Remaudiere and Hennebert (1980) provide strong reasons
to regard both Entomophthora extttalts Hall § Dunn (1957)
and Entomophthora ferruginea Phillips in Houghton and Phil-
lips (1886) as nomtna confusa; we agree with and accept these
determinations. It is impossible to know exactly what fun-
gus is described or illustrated by Hall and Dunn (1957); this
Situation is not eased by the fact that no type was declared
for E. exttralts (nor did the ICBN require typification until
the year after the publication) of ‘this species) or thatethe
cultures deposited by the authors of this taxon in various
major culture collections are found to be either Contdtobolus

513

thrombotdes Drechsler (= Entomophthora virulenta Hall § Dunn)
or Erynta radicans (see Remaudiére and Hennebert 1980). The
fungus regarded to be EF. exttialts by Gustafsson (1965) was
redescribed by Remaudiére and Hennebert (1980) as Erynta
nouryt. Similarly, the real identity of Entomophthora fer-
rugtnea remains indeterminable in view of the three separate
senses in which this name has been applied (see Remaudiére
and Hennebert 1980).

Batko (1964b) offered no explanation for his assignment
of Entomophthora forftculae Giard to Zoophthora. Neither
Remaudiére and Hennebert (1980) nor Remaudiére and Keller
(1980) mentioned this species in their treatments of Zooph-
thora and Erynta, possibly because Giard (1889) mentioned
no secondary conidia (the character used by Remaudiére et al.
to separate these two genera), and possibly because of the
uncertainty which surrounds the identity of this fungus.
PhemconlulamoOteGlardusmspeC1eSma tem exe OuLmuupet Onc om Ga in,
with a length/width ratio of = 3; the conidia in the only
othexrscollections attributed) to this species: are 18=21)x
8-10 Um (Rostrup 1893) and 20-30 x 15-18 um (Petch 1944),
with L/W ratios of € 2. Giard, who did not recognize the
widely used distinction between Empusa (with simple conidio-
phores) and Entomophthora (with branched conidiophores) ,
noted the conidiophores of Entomophthora forftculae to be
only infrequently branched (''peu ramifies''); those of the
fungi reported by Rostrup and Petch could not have been pro-
minently or digitately branched since Petch (1944) reassigned
this species to Hmpusa. Rhizoids were not noted in any of
these three collections (and note that Batko 1964a required
rhizoids to be present in species of Zoophthora!). Despite
the fact that L/W ratios of 3 or more are known only from
Erynta species (with the sole exception of Balazy's undescri-
bed species noted by Humber 198la), the absence of informa-
tion regarding the wall structure and nuclear number of the
Conidid.sand of the actual nature of the yconidiophores:sug-
gests that no definitive generic assignment for Entomophthora
forfteulae is advisable in the absence of other characters
so frequently noted in species of Erynta, It seems doubtful
thatethe collections of fungi by Rostrup (1893) and Peteh
(1944) are conspecific with EF. forftculae Giard; the true
identity of each of these fungi remains to be established by
re-examination of any existing herbarium specimens.

At least two aspects of this emendation of Erynta de-
serve further comment. It is important to re-emphasize that
this nomenclaturally required change from Zoophthora to Ery-
nia has also necessitated the shift of the type species from

514

the very common Erynta radtcans (= Entomophthora sphaero-
sperma Fres. sensu Thaxter) (see Batko 1964a; Remaudiére and
Hennebert 1980) to the less well known Zrynta ovtspora. Re-
gardless of its nomenclator, the fungus which is still most
widely known as Entomophthora sphaerosperma has served as the
type for a genus distinguished primarily by the presence of
rhizoids and branched conidiophores ever since Nowakowski
(1883) affirmed a real taxonomic difference between kmpusa
and Entomophthora.

Entomophthora aphtdis Hoffm. in Fres. was originally
described only from its brown, roughened resting spores; the
conidial state of this species was not found or described by
Hoffmann. In his influential monograph, Thaxter (1888) ap-
plied the name Empusa (Entomophthora) aphtdts to an excep-
tionally common fungal pathogen of aphids which produces
conidia and whose resting spores may be assumed to exist but
have never been found. Remaudiére and Hennebert (1980) were
successful in rediscovering Hoffmann's fungus and demonstra-
ted unequivocally that Thaxter had misapplied Hoffmann's spe-
cific name: The conidia of #. aphtdts Hoffm. are elongate-
fusoid and produce secondary amygdaliform capilliconidia atop
Capillary ‘conidiophoreés. Ihe) presence of capil liconidia in
EH. aphtdis but not in Thaxter's fungus was used by Remaudiére
and Hennebert to place these two fungi into separate genera
as Zoophthora aphtdits (Hoffm. in Fres.) Batko and Erynta neo-
aphtdts Rem. §& Henneb. (the new name provided for Thaxter's
misapplied sense of FE. aphtdis). The emended sense of Erynta
provided here places both of these species together in the
Same genus. » It wil lebevespecially amportanty then, storsstus
dents of these fungi not to confuse these two similar speci-
fic names and to be certain that they do not apply the name
Erynta aphtdts (a very uncommon fungus with a seemingly nar-
row host range and localized distribution) when the fungus
they refer to is actually Erynta neoaphtdts (a very common
Species with a wide host range and occurring in most parts
of the world).

In addition to the species of Hrynta discussed above,
we are aware of two additional new species awaiting publi-
cation: Kramer (1981) is proposing a new species of Hrynta
from snipe flies (Diptera: Rhagionidae). Ben-Ze'ev and Ken-
neth (1981b) are proposing a new species of Zoophthora which
affects froghoppers (Homoptera: Cercopidae); this species
will have to be transferred to Erynta after its formal pub-
lication.

ets)

ACKNOWLEDGEMENTS

Wemwis mC Omtlianicelrs eR So stSODeEL Rh er at korn a andeD:
E. McCabe for their review and constructive comment on this
Manuscripts anders. .o. Kelier, RR: G. Kenneth, and J. PL.
Kramer for providing invaluable information, some of which
Hsm@ettnereiny presspOreinepreparationetor publications

REFERENCES CITED

BATKO, A. 1964a. On the new genera: Zoophthora gen. nov., Trtplospo-
ytum (Thaxter) gen. nov. and Entomophaga gen. nov. (Phycomycetes:
Entomophthoraceae). “Bull. Polon, Acad. Set., Sér. Sci. Biol. 12:
323-326.

BATKO, A. 1964b. Some new combinations in the fungus family Entomoph-
thoraceae (Phycomycetes). Bull. Polon. Acad. Set., Sér. Set. Biol.
12: 403-406.

BATKO, A. 1966a. A new aphidicolous fungus from Poland — Zoophthora
phallotdes sp. nov. Acta Mycol. 2: 7-13.

BATKO, A. 1966b. On the busgenera of the fungus genus Zoophthora
Batko 1964 (Entomophthoraceae). Acta Mycol. 2: 15-21.

BATKO, A. 1974. Filogeneza a struktury taksonomiczne Entomophthoraceae.
In Ewolucja btologtcana: Szktce teorettczne t metodologtczne (C.
Nowinskiego, ed.). Polska Akad. Nauk, Inst. Filozof. Socjol., Wroclaw.

BEN-ZE'EV, I. 1980. Systematics of entomopathogenic fungi of the
"sphaerosperma group" (Zygomycetes: Entomophthoraceae) and their pro-
SPeci com Oluceuineol10lOvUCalepest CONCTOL. sun. Dem Gissentdation.
Hebrew University of Jerusalem.

BEN-ZE'EV, I., and R. G. KENNETH. 198la. JZoophthora ortentaltis sp. nov.,
a fungal pathogen of Aphis cttrtcola (Homoptera: Aphididae), and two
new combinations of other species of Entomophthoraceae. Phytopaga-
eevee 92) 55-42)

BEN-ZE'EV, I., and R. G. KENNETH. 1981b. JZoophthora radtcans and
Zoophthora petcht sp. nov. (Zygomycetes: Entomopnthorales), two spe-
cies of the "sphaerosperma group" attacking leaf-hoppers and frog-
hoppers (Homoptera). Entomophaga 26: 131-142.

BEN-ZE'EV, I., and R. G. KENNETH. 1982a. Features-criteria of taxonomic
value in the Entomophthorales: I. A revision of the Batkoan classifi-

- cation. In preparation for Mycotaxon.

BEN-ZE'EV, I., and R. S. KENNETH. 1982b. Features-criteria of taxonomic
value in the Entomophthorales: II. A revision of the genus Hrynta
Nowakowski 1881 (= Zoophthora Batko 1964). In preparation for
Mycotaxon.

DESCALS, E., J. WEBSTER, M. LADLE, and J. A. B. BASS. 1981. Variations
in asexual reproduction in species of Entomophthora on aquatic insects.
Dane eer. Lee Yi COL.820C,) (728807102.

GIARD, A. 1888. Sur quelques entomophthorées. Bull. Set. France Bel-
gtque 19: 298-309.

GIARD, A. 1889. Sur quelques types remarquables de champignons entomo-
phytes. Bull. Set. France Belgique 20: 197-224.

GUSTAFSSON, M. 1865. On species of the genus Entomophthora Fres. in
Sweden. I. Classification and distribution. Lantbrukshogskol. Ann.
61:8 103-2127,

HALL, I. M., and P. H. DUNN. 1957. Entomophthorous fungi parasitic on
the spotted alfalfa aphid. Hilgardta 27: 159-181.

516

HOUGHTON, W., and W. PHILLIPS. 1886. On Aphte rwmicts, Linn., as a
» pest on the Mangel-Wurzel crops in Shropshire in the autumn of 1885,
and on a fungus destructive of the same Aphis. Ann. Mag. Nat. Htst.
Sera o, Lore lay.

HUMBER, R. A. 198la. An alternative view of certain taxonomic criteria
used in the Entomophthorales (Zygomycetes). Mycotaxon 18: 191-240.

HUMBER, R. A. 1981b. Erynta (Zygomycetes: Entomophthorales): valida-
tions and new species. Mycotaxon 18: 471-480.

HUMBER, R. A. 1982. Strongwellsea vs. Erynta: the case for a phylo-
genetic classification of the Entomophthorales (Zygomycetes). In
preparation for Mycotaxon.

JACZEWSKI, A. A., and P. A. JACZEWSKI. 1931. Opredelitel grtbov,
vob) b, Leningrad:

KRAMER, J. P. 1981. A mycosis of the blood-sucking snipe-fly Sympho-
romyta hitrta caused by Erynta tthacensts sp. n. (Entomophthoraceae) .
Mycopathologta: in press.

MacLEOD, D. M., and E. MULLER-KOGLER. 1970. Insect pathogens: species
originally described from their resting spores mostly as Tartchtum
species (Entomophthorales: Entomophthoraceae). Mycologia 62: 33-66.

MOLLIARD, M. 1918. Sur la vie saprophytique d'une Entomophthora (E.
henrict n. sp.). Compt. Rend. Hebd. Acad. Set. Parts 167: 958-960.

NOWAKOWSKI, L. 1881. O grupie owadomorkéw (Empusaceae). Datenn. III
Zjazdu Lek Przyr. Polak. Krakow., Seke. Bot. 6: 67-68.

NOWAKOWSKI, L. 1883. Entomophthoreae. Przyczynek do znajomésci pasor-
zytnych gryzbkow sprawiajacych pomor owadéw. Pamtetn. Wydz. Akad.
Unteg. w Kradow. 8: 153-183.

PETCH,= I. 1932.) A list) of the entomogenous (fungivot Greate Brigain:
TONS MET me MUCOL MRCOG it? LOL 18s

PETCH, T. 1944. Notes on entomogenous fungi. Trans. Brit. Mycol.
plete, VAS “oalestels) 4 ;

RAUNKIAER, C. 1892. Et Par nye Snyltesvampe. Bot. Tidsskr. 18: 108-111.

REMAUDIERE, G., and G. L. HENNEBERT. 1980. Révision systématique de
Entomophthora aphtdits Hoffm. in Fres. Description de deux nouveaux
pathogenés d'aphides. Mycotaxon 11: 269-321.

REMAUDIERE, G., and S. KELLER. 1980. Reconsidération systématique des
genres d'Entomophthoraceae 4 potentialité entomopathogénes. Myco-
taxon 11: 323-338.

ROSTRUP, 0. 1893. Medd. naturf. Foren. (Copenhagen), p. 87,

ROZSYPAL, J. 1961 Prispevek k biologickému boji proti skudcum obili-
novi parasit larev hrbace osenniho. Sborn. Ceskosl. Akad. Zemed. 24:
85-94.

THAXTER, R. 1888. The Entomophthoreae of the United states) Mem:
BOSCOn SOC NA ws vet.) 4.) 55-201

TURIAN, G. 1957. Entomo-mycoses dans la région de Genéve.. Mitt.
Sehwetz. Entomol. Ges. 30: 93-98.

WATERHOUSE, G. 1975. Key to the species of Entomophthora Fres. Bull.
Brit. Myeol. (50C., 9: loreal.

MYCOTAXON

VO ett eeN On opp 5 1 5.74 October-December 1981
eee ae ee ee oe ee ee Se EE a ee ee ee

REVUE DES LIVRES

par

G.L. HENNEBERT
Book Review Editor, Croix du Sud 3, B-1348 Louvain-la-Neuve
Belgique

THE COELOMYCETES, Fungi imperfecti with pycnidia, acervuli and
stromata, by Brian SUTTON, 696 p., 397 fig., 8°, hard cover, 1980.
Commonwealth Mycological Institute publication, Commonwealth Agri-
cultural Bureaux, Farnham, Slough SL2 3BN, England. Price UK

£920 .005abroadss 133.60.

The Coelomycetes by Sutton is the first comprehensive and modern
contribution on that important wide spread group of fungi. After numerous
publications on the Coelomycetes, including seven issues in the CMI
Mycological Papers and the fundamental and synthetic chapter on Coelomy-
cetes in The Fungi IVA (1973), we are really thankful for such a fine
monograph covering 375 genera and 750 species.

The inestimable progress realized by Dr Sutton in the taxonomy
of these fungi is the recognition of conidiogenesis as a basic taxonomical
criterion to be used at the level of classes and orders, the conidiomatal
structure (pycnidia, acervuli, sporodochia, etc.) and morphology being
useful as a secondary criterion at the suborder level or lower. Although,
in that concept of the taxonomy of conidial fungi, Hyphomycetes and
Coelomycetes would ideally be and are close to being unified and classi-
fied according to the conidiogenesis, the conidioma from the pycnidium to
the obsolete exposed conidial cell representing a continuum, the author
still uses the term Coelomycete here for the practical reason of deli-
miting a workable field. Accepting the major modes of conidiogenesis as
defined at the lst Kananaskis Conference (1969), Sutton proposes two
classes in the Deuteromycotina, the Thallodeuteromycetes and the Blasto-
deuteromycetes, and five orders, the Thallales, Enterothallales,
Blastales, Phialidales and Tretales, instead of the traditional Sphaerop-
sidales, Melanconiales and Moniliales. The structure of the conidioma
' delimits eventually the suborders, f.i., in the Blastales, the Blasto-
hyphineae (blastic Moniliales), the Blastopycnidineae (blastic pycnidial),
the Blastopycnothyrineae (blastic Pycnothyriales) and the Blastostroma-
tineae (blastic Melanconiales).

Notice that two orders, the Enterothallales and the Tretales, are
not reprensented amongst the coelomycetous fungi so far but in the Hypho-
mycetes only. The Coelomycetes traditionally classified in the Pycno-
thyriales and now representing the Thallopycnothyrineae, the Blastopycno-
thyrineae and the Phialopycnothyrineae, are not treated for the simple
reason that almost no information on their conidiogenesis is available.

Sutton discuss the hypothesis of Morgan-Jones, Nag-Raj and Kendrick
(1972) that a disinction might exist between an annellidic conidiogenous
cell and percurrently proliferating phialide but is not convinced . He
prefers to base the distinction between annellidic and phialidic

518

percurrently proliferating conidiogenous cell on the single or repetitive
conidium production by each of the successive cells. There is the reason
why some fungi having conidiogenous cells looking like annellides are
classified as repetitively proliferating phialidic fungi.

In regard of the amount of new information to be included, the
author chose with reason to avoid any already published data and to refer
to good descriptions, illustrations or comments published elsewhere when
possible. Quite a number of genera are therefore not illustrated. But
it would have been of great help to the users to reproduce, at least,
an illustration of those genera, to give at oncea full picture of the
group.The text of the book is concise but clear. The paper, rather thin
but of good quality and flexible, makes the size of the book reasonable
and its manipulation easy. The book is completed with adequate indices
and a glossary.

There is no need to say that the Coelomycetes are economically im-
portant plant pathogens and material deteriogens. Not only should any
mycologist welcome this major contribution to the systematics of fungi,
so also should any plant pathologist, forest manager or microbiologist
concerned with soil biology or material degradation be aware of it.

Furthermore, taxonomists will appreciate being stimulated in their
investigations by an author who, at almost every page of his book, points
out some ways of progress.

A REVISION OF CHRYSOSPORIUM AND ALLIED GENERA, by C.A.N. van OOR-
SHOTS studiessin Mycology n = 20,690 =pann JON toon) amon eens De
cover, 1980. CBS Publicaticon,..Baarn Nederlands PricesHrin Z5,—-

Chrysosporium Corda and the related genera Myceliophthora, Emmonsia,
Zymonema, Trichosporiella, Blastomyces, Glenosporella and Geomyces are
revised, including 38 species and varieties. The redescribed anamorphs
have either thallic or blastic conidiation and rhexolytic dehiscence.
Teleomorphs belong to the Gymnoascaceae, Onygenaceae, Ascosphaeraceae °
and Sordariaceae. The genus Chrysosporium Corda is quite an homogenous
grouping now, Sporotrichum pannorum Link having a distinct position in
Geomyces.

COELOMYCETES. VII. STEGONSPORIUM, by K.T. Van WARMELO and B.C.
SUTION, Mycological’ Papers n- 145, 46 p. 7913) pl... °8 =, paper) covers
1981. Commonwealth Mycological Institute, Kew Surrey, England.
CABLPublicabironasericesf.3).00.

The original spelling Stegonsporium from Corda in Opiz 1926 is
accepted instead of Fries's spelling Stegonosporium 1849. Two species
are maintained, S. pyriforme and S. acerinum. Sixty five species are ex-
cluded or questioned. The excluded species belong to Stigmina, Myxocyclus,
Coryneum, Camarosporium, Trimmatostoma, Dictyodesmium, Neohendersonia and
Camaropellum, with five new combinations. Two new genera, Stegonsporiopsis
and Kaleidosporium are proposed with two new combinations. Stegonsporium
produces annellidic conidiophores with dictyoseptate conidia amongst
filiform paraphyses.

CONTRIBUTION TOWARDS A RATIONAL ARRANGEMENT OF THE CLATHRACEAE,
by D.M. DRING;, 96 p., 2/7 ifie=, 8 . paper cover, 1981, srepranted
from Kew Bulletin, 35(1), 1980. Royal Botanic Gardens, Kew, England.
Price Loe Oor

These are the conclusions of a long-term interest and research in
the Clathraceae by the late Dr. Dring (f 1978) and of his personal

observations in West Africa. It has been fortunate that the unfinished
manuscript could be completed from the author's large collection of notes
and drawings. Dring recognized 8 genera, Azeroé, Blumenavia, Clathrus,
Colus, Ideodictyon, Laternea, Lysurus and Pseudocolus, and 36 species.
These are fully described and illustrated, with synonymies and references
to other valuable illustrations. The specimens examined are all cited,
demonstrating the often restricted geographical distribution. The genus
Clathrus, with 16 species, is the largest one and appears to the author
as the ancester of a diversifying evolution. All the species of the
Clathraceae, the exception of Clathrus ruber and C. hirudinosus repre-
sented in Europe, West Asia and North Africa, are known from the other
parts of the world and many are from tropical areas.

THE GENERA OF FUNGI SPORULATING IN PURE CULTURE, by J.A. von ARX,
Chardeedi tion. s424e peels pie oS ee kardacovers =| 96170 mCramer:
Fl 9490 Vaduz, Lichtenstein. Price DM 120.-, for subscribers 100.-.

This third edition of Dr von Arx's book is a fully revised and ex-
panded version of the previous ones. The classification of the fungi has
been emended according to recent advances. The Dothideales include the
Myriangiales, Dothiorales and Pseudosphaeriales. The Pezizales include
the Tuberales. The Tilletiales are reintroduced. The Ustomycetes (Ustila-
Sinales, Exobasidiales, Taphrinales, Sporobolomycetales and basidiomy-
cetous yeasts) are distinguished from the Endomycetales on morphologi-
cal grounds and wall composition. Basidiomycetes are not otherwise trea-
ted. All together 853 genera are accepted, of which only 370 are given
an illustration, one third of these illustrations being new and based on
Yecent studies. About 50 genera, some of them being recently described,
are reduced into synonymy. Some genera rejected in recent years are re-
introduced, e.g. Endomycopsella, Karakulina and Bipolaris. Illustrations
and dichotomous keys result from studies of the fungi in pure culture,
independent of their natural substrate. Keys have been adjusted to up-
dated terminology. The 1100 literature titles include the most recent
ones published in 1980. This edition, certainly like the previous ones,
will be appreciated by those who have to identify fungi from sporulating
cultures.

MARINE MYCOLOGY, The Higher Fungi, par Jan KOHLMEYER et Erika
KOHLMEVER (t210).090 tpe, 1270 fie, eo a erelaeyto1lé (1979.
Academic Press, N.Y. Price US $ 71.50.

149 Ascomycétes, 4 Basidiomycétes, 56 Deuteromycétes filamenteux
et 177 levures sont les champignons supérieurs récoltés 4 ce jour prés
de et dans la mer. Des 209 espéces filamenteuses rencontrées, 191 sont
spécifiques au milieu marin. C'est dire qu'un ouvrage intitulé Marine
Mycology ne saurait @étre de pure taxonomie sans aussi considérer 1'éco-
logie. C'est cette alliance qu'ont fort bien réussie les deux auteurs.
Aprés un exposé des méthodes (chap. 2) 1'écologie des champignons cétiers
et marins met en évidence les relations champignon-substrat chez les
espéces de pleine eau, d'estuaires, de sables, des écumes, des marais
salés, des algues , des lichens, des mangroves et des débris animaux et
végétaux (chap. 3-19 et 21). Dans les chapitres touchant la taxonomie
(chap. 20, 22-28; 421 p.) les auteurs proposent d'abord des clés dicho-
tomiques basées sur la morphologie sporale ou conidienne et finement il-
lustrées par Erika Kohlmeyer. Le principal chapitre 26 comporte la des-
cription taxonomique des 209 champignons filamenteux, avec synonymie,
photographies et bibliographie. Une nouvelle espéce et 7 combinaisons
sont proposées. 49 rejections de noms douteux ou invalides sont énoncées.

5.0

Dans le dernier chapitre,les levures sont répertoriées, sans description
mais avec indication de leur habitat.Prés de 1400 références bibliogra-
phiques complétent l'ouvrage. En mycologues avertis, les auteurs témoi-
gnent dans ce livre d'un sens aigu du détail et ume maniére esthétique
de le mettre en valeur. Ce livre est une "somme" de ce domaine particu-
lier de la mycologie. Sa qualité suscite 1'admiration.

MICROBIAL POLYSACCHARIDES AND POLYSACCHARIDASES, par R.C.W. BERKE-
LEY, G.W. GOODWAY and D.C. ELLWOOD édit., Publication de la Society
for general Microbiology, xvinte4v79'p.,) 211 oe oe relic. ag 19s
Academic Press, N.Y. Prix US $947.00.

Ce livre est la synthése de deux symposiums tenus 4a Aberdeen en
1978, l'un sur The microbial extracellular Polysaccharides du Groupe pour
1'étude des surfaces et membranes de la Cellule microbienne, l'autre sur
The microbial Degradation of Polysaccharides de la Society for general
Microbiology.

Sans doute, cet ouvrage sort du champ habituel de la mycologie.
Cependant la composition différente des parois des champignons suivant
leur classe taxonomique est une raison de considérer cet ouvrage de portée
aussi fondamentale qu'appliquée. La détection, l'analyse et la production
de polysaccharides et des enzymes qui les dégradent sont les objectifs
d'une recherche de plus en plus active. Ce sont surtout les polysaccha-
rides extracellulaires, comme le pullulan de 1'Aureobasidium pullulans,
qui retiennent l'attention. La paroi fongique est donc un organe essen-
tiel. Sa composition, sa structure microscopique fine, sa perméabilité,
la structure biochimique, l'activité et la solubilité de ses polysaccha-
rides sont autant d'aspects traités dans ce livre. La paroi se détruit
autant qu'elle se construit. Les polysaccharidases produites par les
champignons eux-mémes ou les bactéries associées agissent au niveau de
la paroi fongique et dans le milieu, transformant les polysaccharides en
nutrients recupérables. Ainsi les cellulases, chitinases, amylases, glu-
canases, pullulanases des champignons ne sont pas les moins connues.

Ce livre s'adresse non seulement au microbiologiste et biochimiste indus-
triel, mais aussi le mycologue curieux de la nature exacte des champignons.

STEROLOGICAL METHODS. VOL. I. PRACTICAL METHODS FOR BIOLOGICAL
MORPHOMETRY,--par ©aR, WEIBEL, ©515ep. , milli Seretie9 toile, 31979.
Academic Press. Price US $ 66.50.

La premiére approche d'un étre vivant est d'abord d'en saisir la
forme. Au travers du microscope, ot le relief disparait, la forme est
celle d'une image, d'un profil, d'une section. Le microscopiste, le bio-
logiste, le mycologue traduit la vision de la forme (morphoscopie) en un
tracé d'abord (morphographie), en une description ensuite (morphologie),
en une mesure enfin (morphométrie). La promotion récente de la taxonomie
numérique accentue la nécessité d'une morphométrie méthodique et tradui-
sible en termes de réalité. La recherche de méthodes morphométriques
valables est le but delastéréologie. Ces méthodes sont mathématiques et
visent a relier les paramétres tridimensionnels d'une structure spatiale
aux mesures bidimensionnelles des sections de cette structure. Inversé-
ment, elles fournissent la réponse a la question: comment interpréter
l'image vue au microscope comme expression de la réalité spatiale.
L'auteur ne traite pas des fondements mathématiques des méthodes, les
réservant au volume II. Le livre est donc d'un niveau mathématique acces-
sible & qui posséde des notions de mathématique moderne. Par voie
d'exemples, de photos et de graphiques, 1'auteur explique les méthodes
d'échantillonage des structures A mesurer et la réalisation de mesures

ork

qui ne faussent pas la réalité. Il serait intéressant que des mycologues
expérimentent les méthodes morphométriques proposées par la stéréologie
au profit de la taxonomie des champignons.

BULLETIN OF MYCOLOGY. VOL. I. N°l, edited by Sultan AHMAD and S.H.
IQBAL, Department of Botany, University of the Punjab, New Campus,
Lahore, Autumn 1980.

We welcome the appearance of this new journal devoted to taxonomic
mycology from Pakistan, a country where surely much waits to be disco-
vered and described. The first issue of the Bulletin of Mycology contains
interesting papers on freshwater Hyphomycetes, on coprophilous Ascomyce-
tes, on Agaricales and other fungi from Pakistan. We hope that the new
journal receives the attention that it deserves.

SEXUALITY AND PATHOGENICITY OF FUNGI, edited by R. VANBREUSEGHEM
andeCh.s DEWVROEY. 2505p... 111, ,) 8°, hard «cover, 1981. Masson, 120
Bd, Saint Germain, 75280 Paris.

This is the Proceedings of the Third international Colloquium on
Medical Mycology organized by the Prince Leopold Institute of Tropical
Medecine, Antwerp, Belgium.

The question raised at the beginning of the colloquium "Is there a
relationship between sexuality and pathogenicity to man in fungi?" recei-
ved an answer far beyond any hope. That answer is that no relation between
mating type and symptoms has been observed so far, but that a close rela-
tion between the mating type and the host or biotope is demonstrated and
that that relation is determining particular geographical distribution.
Beside the pathogenicity of mating types, serotypes, + and - types, sexual
and asexual spores, many correlative aspects of the pathogenic fungi have
heen treated by the 32 contributors, like taxonomy and the relation ana-
morph-teleomorph, gentics, somatic hybridization, sexual stimulation,
sexual test strains, filamentization of yeasts. The book answers a number
of questions raised in the taxonomy and ecology of the fungi. It offers
valuable readings for any mycologist and for plant pathologists.

YEAST GENETICS AND MOLECULAR BIOLOGY 1980. Reports, by A, GOFFEAU and
J.M. WIAME. 102 p., 8°, paper back, 1980. Louvain-la-Neuve, Lab.
Enzymology, Croix du Sud 1, 1348 Louvain-la-Neuve, Belgium, Price
Workshop reports $ 10, Abstracts book $ 20.

This is the Workshop Reports of the tenth International Yeast Con-
ference held at Louvain-la-Neuve, September 1980. It contains 24 plenary
presentations covering the research progresses on Saccharomyces cerevi-
siae about its chromosome structure and replication, RNA, protein synthe-
sis, cell cycle, sporulation, recombination, mutagenesis, radiation repair,
expression of cloned genes, mating, regulation of C and N, metabolism,
mitonchondria, killers, genetics engineering, evolution and taxonomy.

It certainly leads to conclude that Saccharomyces cerevisiae is a
favorite model for studying the structure and the operation of eukaryo-
tic cells. The Abstracts Book was distributed at the Conference but is
still available, it reproduces the abstracts of each of the 200 posters
presented.

INTRODUCTION TO FUNGI, by John WEBSTER, 2d edition, 669 p., 331 fig.
8°, paper or hard cover, 1980. Cambridge University Press. P.O. Box
110, Cambridge, England. Price h.c. £ 30, p.c.. 9.95.

522

After the publication of the first edition, in 1970, advances have
been made in all fields of mycology. Where necessary the text has been
altered or expanded. The literature cited is twice in number now. The
author adopts the Ainsworth, Sparrow and Sussman's system of the Fungi,
as a separate kingdom. The Myxomycetes are more extensively treated.
The anamorphic fungi, the Deuteromycotina, receive a special chapter
with a description of their conidiogenesis. New and detailed accounts
are added on three ecological groups of fungi, the aquatic, the preda-
cious and the seedborne imperfect fungi. This reinforces the aim of the
author to be not purely taxonomical but to describe the fungi in their
life behaviour. The clarity of Webster's writing, the precision of the
data, the excellent selection of original photographs and drawings
are prominent characteristics of the book. The book is an excellent
manual for the university level. For reaching constantly the present
limits of knowledges, it concerns all students of mycology.

HOW TO KNOW THE TRUE SLIME MOLDS, by M.L. FARR, in Pictured Key
Nature, Series 5132 px loostiec.., 2oxl/ecm spirale binding goLe
Wm. C. Brown Co. Publ., 2460 Kerper Bd., Dubuque, Iowa 52001, USA.

This "pictured key" to the Myxomycetes is intended to be introduc-
tory and practical. The introduction shortly explains what Myxomycetes
are, how they live, where they growth, how to collect them, how to keep
them alive, how to look at them, how to describe them, how to preserve
them. The keys to orders, families, genera and species are all dichotomous
Specific descriptions, comments and illustrations are given at every spe-
cies keyed out. Illustrations are mostly line drawings picturing frui-
ting bodies and, eventually, capillitium and spores. At every taxonomic
level, indeed, the keys are constructed from both macroscopic and micro-
scopic characters, They thus are not to be considered as field keys.

They cover a selection of 276 common North-American species of Myxomycetes.

TAXONOMY OF THE INDIAN MYXOMYCETES, by T.N. LAKHAMPAL and K.G.
MUKERJI.- in Bibliotheca Mycologica vol= 9785 "53l=p.5, 40Mfiee,es a,
hard cover, 1981. J. Cramer, Fl 9490 Vaduz, Lichtenstein. Price:
DMV1205 subscription DM 96.

The authors have realised two aims. In the first part of the book,
they describe, in a taxonomical order, all the species of Myxomycetes
that they recorded in 1000 specimens from their respective collecting
fields, i.e. Himalahal Pradesh and Delhi. The Taxonomy is that of Martin
and Alexopoulos (1969) modified by Alexopoulos in Ainsworth, Sparrow and
Sussman (1973). The illustrations are assembled in 28 full-page plates
of black and white photographs of standard quality and 12 plates of ra-
ther poor line-drawings. In the second part of the book, the authors pro-
duce an updated index of all the Indian Myxomycetes records. disposed in
taxonomical order, with reference to the pertinent literature. The contri-
bution is important. Out of the 450 presently known species of Myxomyce-
tes, 293 are recorded in India and, out of those, 111 are recently recor-
ded, by the authors mostly, after the publication in 1977 of Dr Thind's
monograph.

BIODEGRADATION ET HUMIFICATION, ATLAS ULTRASTRUCTURAL, par G.M.OLAH,
0; REISINGER et*G.) KILBERTUS 33334 0p. 51998 fie.5..4 eo cartonna.s19/6.
Les Presses de l'Université Laval, Québec. Librairie Vuibert, Paris.

—

En 1974, était organisé, a Nancy, le ler Colloque international sur
le théme "Biodégradation et Humification" (Mycotaxon IV(1):317, 1976).

523

Les auteurs, organisateurs de ce Colloque, ont rassemblé en un atlas leurs
documents de microscopie électronique illustrant d'une part les agents de
la biodégradation dans le sol (bactéries, champignons, algues bleues,

et microfaune) et d'autre part les victimes, la matiére organique vivante
en dépérissement (celle des algues, champignons, bryophytes et phanéro-
games). L'atlas est divisé en chapitres introduits par un texte et un
schéma expliquant le processus de la biodégradation dans le sol. Les
illustrations font face a une légende bréve, & mon avis trop bréve, mais
suivie d'un grand nombre de références & la littérature. Les photographies
sont de bonne qualité et suscitent l'intérét par leur choix et leur ori- 1
ginalité. On y voit des vues inattendues de bactéries du sol ou coloni-
sant des hyphes fongiques. Des images en ombrage et en transmission il-
lustrent la sporulation de divers champignons, un asque operculé béant,
des annellides, des conidies en grappe, des pores septaux, la perforation
bactérienne de parois fongiques, la désorganisation enzymatique de celle-
ci, et la repousse interne d'une hyphe de régénération. Cet atlas est
didactique et permettra de visualiser, dans l'enseignement, un des grand
phénoméne de la biologie du sol, le "turn-over" de la matiére organique.

COMPENDIUM OF COTTON DISEASES, par G.M. WATKINS, éd., in The Disea-
SemCOmpendiasSeries pallu, Pomp. 64ers oo Oy pile COL. Ga BLOG Le

The American Phytopathological Society, 3340 PilotKnob Road, St Paul,
Minnesota 55121. Prix US$ ll.

Ce compendium, comme les précédents sur les maladies du soya, du
froment, de la luzerne, du mais, continue une . série fort utile 4 la
clinique des plantes cultivées. Plus de quarante spécialistes ont contri-
bué 4 la rédaction et l'illustration de celui-ci. Prés d'une cinquantaine
de champignons responsables d'une pathogénése fongique du cotonnier
sont envisagés, la symptomatologie et 1'épidémiologie décrites et le
traitement indiqué. A coté des maladies fongiques, les autres maladies
biotiques et abiotiques sont aussi décrites et illustrées. Une clé syn-
thétique des symptOmes permet l'identification rapide des maladies.

THE CORTICIACEAE OF NORTH EUROPE. Vol. 6. PHLEBIA-SARCODONTIA, par
John ERIKSSON, Kurt HJORSTAM et Leif RYVARDEN, p.1051-1276, fig.
DEI -ODShmO wee Droche sl Iolerunes tl ora, er..07 eboxno> se bkindern ,

Os OmS se Norway. Prix gNkresl20.

Sont déja parus et recensés dans Mycotaxon les volumes 2 (Aleuro-
discus - Confertobasidium), 3 (Cornicium - Hyphoderma), 4 (Hyphodermella
- Mycoacia) et 5 (Mycoaciella - Phanerochaete). Dans le présent volume,
les auteurs décrivent les genres Phlebia (31 espéces), Repetobasidium
(8 espéces), Piloderma, Resinicium (3 espéces chacun), Phlebiopsis, Ra-
dulomyces (2 espéces) etPhysodontia, Plicatura, Plicaturopsis, Podoscypha,
Pseudomerulius, Pseudoxenasma, Pulcherricium, Punctularia, Radulodon,
Ramaricium, Repetobasidiellum et Sarcodontia monospécifiques. Les auteurs
proposent 1 nouveau genre (Repetobasidiellum), 9 nouvelles espéces dont
3 en Phlebia, 14 nouvelles combinaisons dont 5 en Phlebia et un nom nou-
veau. Certainement, ce volume est attendu, comme le sont aussi les suivants.

THE FISCHER-SMITH CONTROVERSY: ARE THERE BACTERIAL DISEASES OF
PLANTS?, par Erwin F. SMITH et Alfred FISCHER, introduit et tra-
duit de 1'allemand par C. Lee CAMPBELL, Phytopathological Classics
n° 13, xviii + 65 p., 40 fig., 8°, broché, 1981. The American
Phytopathological Society, St Paul, Minn., USA. Prix US $ 8.50.

Erwin F. Smith, American plant pathologist, USDA, Washington, who
has demonstrated experimentaly the pathogenicity of bacteria to plants

524

refutes theassertion of Alfred Fischer, a German professor in Botany at
the University of Leipzig, of the non-existence of bacterial diseases.

It was in 1897. Fischer answered by unjustifiable critics about the
Smith's observations and experiments. Smith then took his own defence,

ina long and convincing paper (1902). As a result, the existence of
bacterial diseases was accepted, and the rivals were both promoted, the one
as director of the USDA Plant Pathological Laboratory, Washington, the
other as professor and director of the Botanical Gardens at the Universit
ty of Basel.'The facts ardé\God's, the theories‘are human" (Lord Ripon, 1874).

TROPICAL MYCORRHIZA RESEARCH, par Peitsa MIKOLA édit., Oxford
SciencesPublicatsonsiyexiy +2 /ONp., oil Gere lie lL OOOmshOrethe
International Foundation of Science, Oxford University Press,
Walton ‘Street, (Oxford s0xX2,6DP; Englands Prix=e ilo.

Ce volume est constitué des 31 communications présentées a 1'Inter-
national Workshop on Tropical Mycorrhiza tenu au Ghana, en 1978, par 1'
International Science Foundation (IFS). Elles se regroupent en quatre
thémes: les ectomycorhizes en région tropicale, les mycorhizes de la
végétation naturelle, nutrition mycorhizique des plantes tropicales et
mycorhizes des plantes agricoles. Il est intéressant de constater le
role grandissant des ectomycorhizes dans la reforestation des zones tro-
picales dénudées. D.H. Marx fait ume revue trés large des possibilités
d'inoculation de champignons mycorhizogénes, a4 travers le monde, et des
méthodes utilisées. D'autres donnent les résultats particuliers obtenus
en leur pays. La recherche explore aussi les mycorhizes naturelles des
essences forestiéres et des plantes cultivées tropicales, en particulier
de 1'Hevea, de 1'Elaeis, du cacaoyer, des Citrus et des graminées. Si on
recherche les facteurs qui peuvent leur @tre favorables, on s'inquiéte
de plus en plus de l'effet néfaste sur les mycorhizes des fertilisations
chimiques et des herbicides en forét comme en agriculture. La mycorhization
est un phénoméne tel qu'il importe de le connaitre pour le maintien du
monde végétal et du notre.

DEVELOPMENTAL MICROBIOLOGY. par John PEBERDY, in Tertiary Level Bi-
ology Series, 230yp-,,11U., 12° ,apaper back,, 1980... Blackie-& Son Ltd
Bishopbriggs, Glascow G64 2NZ. Prix £ 8.50.

Ce petit livre de microbiologie est axé sur les phénoméne de crois-
sance et de reproduction. Il en montre les aspects les plus fondamentaux
de structure, de biochimie et de génétique cellulaires et les plus appa-
rents de la morphologie, de l'accroissement et de la formation sexuée et
asexuée de spores, avec le souci d'expliquer la relation entre ces aspects.
Si dans un tiers du livre il est question de bactéries et virus vus sous
les mémes aspects, ce livre est particulier par le choix des champignons
comme matériel privilégié d'étude du développement (levures, myxomycétes,
champignons filamenteux, champignons sexuées et asexuées). Ce livre est
destiné au niveau du baccalauréat ou "undergraduate".

BIOLOGICAL MEMBRANES, THEIR STRUCTURE AND FUNCTION, par R. HARRISON
et G.G. LUNT, Tertiary Level Biology Ser«, 2e.ed., 286 po gelll oe toe
L980... Blackiess SonelLtd. «Glas cow. .er isaac 3

La structure de la membrane cellulaire, sa composition biochimique,
les mécanismes fonctionels de reconnaissance des macromolécules et de
leur transport par la membrane cellulaire sont les sujets traités par
les auteurs dans un langage clair et adapté au niveau universitaire.
Cependant nous regrettons que ce livre n'envisage que la cellule animale
etqu'ilnementionne qu'en passant les membranes végétale et fongique.

MYCOTAXON

Onc ar Ome 5 a OO October-December 1981

NE Omi ia Ce eaS

XV PACIFIC SCIENCE CONGRESS

The XV Pacific Science Congress will be held February
1-11, 1983, in Dunedin, New Zealand. The Pacific Sci-
ence Association is responsible for the program, which
is sponsored by the Royal Society of New Zealand. A
fisrt circular may be obtained from The Secretary Gene-
ral, 15th Pacific Science Congress, P.O. Box 6063, Dune-
din North, New Zealand.

IMC: ON HOUSING COSTS IN JAPAN

Though it is indeed true that Japan is currently a
very expensive place to visit, the organizing committee

of IMC3 is well aware that most mycologists will not be
able to spend great sums of money on housing for the
congress. Dr. Keisuke Tubaki, a member of the Mycolo-

gical Society of America's ad hoc committee on IMC3 ar-
rangements, has assured us that though the meetings
are planned for a downtown Tokyo hotel-convention faci-
lity, there will be many hotel rooms available close by
in fess prestigious hotels in the $15-20/night (1981
prices) range. There have apparently been rumors that
housing costs would be substantially higher. Our Japa-

nese hosts will do all they can to keep costs down for
participants. The same concerns will be kep in mind
for pre-congress and post-congress tours, workshops,
and field trips. The Tokyo meetings will be from 28

August through 3 September, 1983.

526

AUTHOR INDEX, VOLUME THIRTEEN

ALCORN, J. L. Cochliobolus ravenelit sp. nov. and C. trtpogonts sp. nov.
339-345

ALCORN, J. L. Ascus structure and function in Cochltobolus species 349-
360

ARENDHOLZ, WOLF-RUDIGER & RICHARD P. KORF. The publication date of Arend-
holz's thesis on leaf-inhabiting Helotiales 187-190

BAKER, JOHN G., IRA F. SALKIN, DAVID H. PINCUS & RICHARD F. D'AMATO. Can-
dida paratroptealis, a new species of Candida 115-119

BALAZY, STANISLAW, see MIETKIEWSKI & al.

BEN-ZE'EV, ISRAEL, seé@ HUMBER §& BEN-ZE'EV

BORELLI, DANTE, see McGINNIS §& BORELLI

CANDOUSSAU, FRANCOISE, see PFISTER §& CANDOUSSAU

CRANE, J. L., see HEWINGS & CRANE

D'AMATO, RICHARD F., see BAKER & al.

DUMONT, KENT P. Leotiaceae III. Notes on selected temperate species re-
ferred to Helottum and Hymenoscyphus 59-84

GAMS, W. §& V. HOLUBOVA-JECHOVA. Chlortdtwn and some other dematiaceous
hyphomycetes growing on decaying wood. Corrections and additions 257-258

GOCHENAUR, S. E. Cyrenella elegans gen. et sp. nov., a dikaryotic ana-
morph 267-277

GRUFF, SUSAN C., see KORF & GRUFF

GUZMAN, GASTON, see SCHROEDER §& GUZMAN

HENNEBERT, G. L., Revue des Livres 278-286, 517-524

HENSSEN, A. & B. RENNER. Studies in the lichen genus Psoroma 1: Psoroma
tenue and Psoroma etnnamomeum 433-449

HEWINGS, ADRIANNA D. & J. L. CRANE. The genus Codinaea. Three new species
from the Americas 419-427

HJORTSTAM, KURT. Notes on Corticiaceae (Basidiomycetes). VIII. Two new
species of Tubultecrtnts 120-123

HJORTSTAM, KURT. Notes on Corticiaceae (Basidiomycetes). IX. Three new
combinations in Hypochnitctellum 124-126

HJORTSTAM, KURT §& LEIF RYVARDEN. Studies in tropical Corticiaceae (Basi-
diomycetes) III. Two new species of Laxttextuwmn 35-40

HOLUBOVA-JECHOVA, V., see GAMS §& HOLUBOVA-JECHOVA

HUMBER, RICHARD A. An alternative view of certain taxonomic criteria used
in the Entomophthorales (Zygomycetes) 191-240

HUMBER, RICHARD A. Erynta (Zygomycetes: Entomophthorales): validations
and new species 471-480

HUMBER, RICHARD A. & ISRAEL BEN-ZE'EV. Erynta (Zygomycetes: Entomophthor-
ales): emendation, synonymy, and transfers 506-516

JAHNS, H. MARTIN. The genus PtZophorus 289-330

JENKINS, DAVID T. A new species of Amantta 112-114

KANE, JULIUS, IRA F. SALKIN, IRENE WEITZMAN & CATHERINE SMITKA. Trtcho-
phyton raubitschekit, sp. nov. 259-266 .

KOHN, LINDA M. A preliminary discomycete flora of Macaronesja: Part 3,
Hyaloscyphaceae subf. Trichoscyphelloideae 145-149

KOHN, LINDA M. Sclerotinia bresadolae Rick, a taxonomic synonym of Ctbo-
rtnta candolleana (Lév.) Whetzel 405-406

KORF, RICHARD P. Marcelle Le Gal, a reminiscence 1-4

KORF, RICHARD P. A preliminary discomycete flora of Macaronesia: Part 2,
Hyaloscyphaceae subf. Arachnopezizoideae 137-144

KORF, RICHARD P. A preliminary discomycete flora of Macaronesia: Part 6,
Geoglossaceae 361-366

KORF, RICHARD P. & SUSAN C. GRUFF. Discomycetes Exsiccati, fasc. IV 5-15

KORF, RICHARD P., see ARENDHOLZ & KORF

LEGER, J. C. Les Hymenochaete a éléments hyméniens pinnatifides 241-256

LOWY, B. A new species of Daeryoptnax from Brazil 428-430

LUNGHINI, D., see ONOFRI & al.

LUSTRATI, L., see ONOFRI & al.

McGINNIS, MICHAEL R. §& DANTE BORELLI. Cladosporiwn banttanum and its
synonym Cladosportum trtchotdes 127-136

MIETKIEWSKI, RYSZARD, RICHARD S. SOPER §& STANISLAW BALAZY. Notes on Zo-
ophthora oeetdentalts (Thaxter) Batko (Entomophthorales: Entomophthor-
aceae) 41-49

NAKASONE, K. K. Cultural studies on Porta etnerascens, P. rivulosa, and
P. subvermtspora (aphyllophorales, Basidiomycotina) 105-111

OLIVER, LOLA K. & KEITH VAN CLEVE. Notes on soil fungi isolated from a
15-year-old aspen stand in interior Alaska 369-372

ONOFRI, S., D. LUNGHINI, A. RAMBELLI & L. LUSTRATI. New dematiaceous hy-
phomycetes from tropical rainforest litter 331-338

PALM, MARY E. & ELWIN L. STEWART. Pithomyces pavgti, a new combination
for Trichocladtum pavgtt and Pithomyces funtculosa 465-468

PFISTER, DONALD H. & FRANCOISE CANDOUSSAU. The psilopezioid fungi. VII.
A new species of Pstlopezta from France 367-368

PFISTER, DONALD H. §& FRANCOISE CANDOUSSAU. The psilopezioid fungi. VIII.
Additions to the genus Pachyella 457-464

PINCUS, DAVID H., see BAKER & al.

RAMBELLI, A., see ONOFRI & al.

REDHEAD, S. A. §& R. SINGER. Restnomycena gen. nov. (Agaricales), an ally
of Hydropus, Mycena and Baeospora 150-170

REDHEAD, S. A. §& J. A. TRAQUAIR. Coprtnus sect. Herbicolae from Canada,
notes on extralimital taxa, and the taxonomic position of a low tempera-
ture basidiomycete forage crop pathogen from western Canada 373-404

RENNER, B., see HENSSEN §& RENNER

RYVARDEN, LEIF. Type studies in the Polyporaceae 13. Species described by
J. H. Léveillé 175-186

RYVARDEN, LEIF, see HJORTSTAM §& RYVARDEN

SALKIN, IRA F., see BAKER & al., see KANE & al.

SCHROEDER, ROBERT F. § GASTON GUZMAN. A psychotropic fungus in Nepal 346-
348

SIMMONS, EMORY G. Alternaria themes and variations 16-34

SIMMONS, EMORY G. dHalystomyces, a new dematiaceous genus from Arizona's
Sonoran Desert 407-411

SINGER, R., Seé@ REDHEAD §& SINGER

SMITKA, CATHERINE, see KANE @ al.

SOPER, RICHARD S. New cicada pathogens: Massospora cteadettae from Au-
stralia and Massospora pahartae from Afghanistan 50-58

SOPER, RICHARD S., see MIETKIEWSKI @ al.

STEWART, ELWIN L., see PALM §& STEWART

TAVARES, ISABELLE I. Validation of the Herpomycetineae and Herpomycetace-
ae in the Laboulbeniales 469-470

TERADA, KATSUYUKI. Osortomyces, a new genus of the Laboulbeniales from
Taiwan 412-418

TOYAZAKI, NORIHIRO. An undescribed pleomorphic species of Codinaea 450-
456

TRAQUAIR, J. A., see REDHEAD §& TRAQUAIR

TUCKER, BRUCE E. A review of the nonentomogenous Entomophthorales 481-505

VAN CLEVE, KEITH, see OLIVER §& VAN CLEVE

WEBER, WILLIAM A. Lichenes Exsiccati distributed by the University of Co-
lorado Museum, Boulder: Fascicles 1-15, Nos. 1-600, 1961-1979 85-104

WEITZMAN, IRENE, see KANE & al.

ZANG, MU. Sinotermttomyces, a new genus of Amanitaceae from Yunnan, China
171-174

528

INDEX TO FUNGOUS AND LICHEN TAXA, VOLUME THIRTEEN

This index includes genera,
specific taxa. New Taxa are
they are published.

itself anh index, reference
HEX

Acarospora see 86-101

Acaulopage 276

Actinomyces 370

Agaricus decurrens 156; rhododendri
155, WSS Uiskikelkkeolitel sists}, silo

Agrocybe tibetensis 347

Alectoria see 86-101

Aleuria annamitica 458

IMEC WO, iWsh AO, 245 Wh, 7S, 74).

150-

327 407 oe GiheehOSsthatac we noltinicantes
20; '‘gr' Erostratae, Solitariae 16; al-
ternata 26, 29, 30; chlamydospora 16,
iS, 24, 282 CaevySselaiiaeiml iO, ws, wes

helianthi 16, 18, 19; limaciformis 16,

24, 25; longipes 16, 28-33; molesta 16,
17-19; mouchaccae 16, 18, 20, 21; ra-
dicina 16; solani 28; tenuis 30; tenu-

issima 16, 26, 29, 30;
27; zinniae 16

Ama miliice a ill Zemin tesSeGL
114; subsect Solitariae 112, 114; stirps
Polypyramis 114; stirps Strobiliformis
114; stirps Virgineoides 114; marginata
112-114; muscaria 347

Amauroderma schomburgkii 181

Amyloathelia 124

Anaptychia see 86-101;
multifida f circinalis

Herwienine Wea, AS,

Eepiidellia ws! 12;

leucomelaena v
101; f verruci-

fera 102; podocarpa v stellata 102
Ancylistes 201, 210, 481, 483-487, 489,
492, 493, 501, 502; closterii 482, 484-
486) 488") 490, 49Sn OOS ee netrite 401
482, 484-486, 488, 490, 493, 503; pfeif-

feri 482, 484, 488, 490, 493, 503
Anthracobia macrocystis 7
Anthracothecium ochraceoflavum 86

Anzia see 86-101

Arachnopeziza 6, 137; '‘gr' Anomalae 14;
‘gr' Typicae 14; aranea f. aranea
138; f. monilipila 138, 139; aurata
Uv, UG, WS eile 1S, Wests, Wekale

Cangido=fUilVagOuss lS:au conmUtamsO alo.
14; leonina 6, 13; obtusipila 138, 141,

142; trabinelloides 6, 14; zonulata
WSteh. RAID. aie

Arachnoscypha 139; aranea 138

Arctomia fruticosa 98

Arthonia see 86-101

Arthopyrenia halodytes 87

Arthothelium see 86-101

Arthroderma benhamiae 260, 261; simii
260, 261

Aureobasidium 407

Bacidia see 86-101

Baeomyces see 86-101, 328; acicularis 298
Baeospora 9150505 151-95 168: 3 169 pallida
LO loG

infrageneric taxa,

in boldface as are the pages where
The Lichenes Exsiccati
to which is

species, and infra-
paper (pp. 85-104) is

indicated by the notation

Ballocephala 200, 204, 210, 481, 483-489,
491-493, 498, 501, 502; pedicellata
482, 484, 485, 488,, 490, 493, 503;
sphaerospora 482, 484-486, 488, 490,
493, 503; verrucospora 482, 484-486,
488, 490, 493, 498, 503

Basidiobolus 195, 198, 201, 204, 206-208,
210, 481, 487, 489, 501, 502; -micro=
sporus 195, 208, 489

Belonia americana 87

Beltrania mangiferae 335; muelleri 335;

onirica 331, 333-335, 337; querna 335
Bipolaris 339, 344; micropus 349; rave-
nelii 340, 341, 345; tripogonis 344

Blastocladia pringsheimii 275

Botryobolus parasiticus 482

Boudiera dennisii 5, 7

Bryoria see 86-101

Buellia see 86-101; straminea 103

Calicium see 86-101

Caloplaca see 86-101

Caloplacopsis submexicana 88

Candelariella see 86-101

Candelina submexicana 88

Candida 115; paratropicalis 115-118; stell-
latoidea 117; tropicalis 116, 117

Catenaria auxiliaris 483

Catillaria see 86-101

Cavernularia lophyrea 88

Cenomyce acicularis 298

Cetraria see 86-101

Cetrelia chicitae 88

Chaenotheca brunneola 88

Chaetopsina 333

Chaetosphaerella 257

Chaetosphaeria 450; callimorpha 426;
dingleyae 425; fusiformis 257; fusi-
spora 257; talbotii 426

Chiodecton see 86-101

Chloridium 257, 450; codinaeoides 450,
455

Chondropsis semiviridis 88

Chryseidea 331, 333; africana 331-333,
SG

Chrysosporium pannorum 371

Ciboria 79

Ciborinia 405, 406; candolleana 405, 406

Cladia see 86-101; galapagosensis 101;
polia 101

Cladina galapagosensis 85, 89, 101; po-
lia 85, 89, 101; subtenuis 89

Cladonia see 86-101; pileata 328

Cladosporium 127, 407; bantianum 127-
130 01325 eee 4 ee SORT Ghoncdesmen| 2a.
W285 S2 ela hos

Clavaria nigrita 362
Clavariana 276

Clavatospora 276

CoelhMmi@nelwis Ss SYA, sya GYAG) eit).
35050953 00-358 NDI Colom 5350: carbo—
num 350, 351; cymbopogonis 344, 349,
35), sd, A, esses Cwineclomvls
350, 351; geniculatus 350, 351; hawai-
iensis 344, 353, 356; theterostrophus
34953515, 352, 3545" homomorphus 349—
35 le intermedius 350, 351; kusanoi
Syl. S285 Sse, See VWMMEUS SEO, Se)e
miakei 350; nodulosus 349-351; palmi-
vora 350; ravenelii 339, 341, 342,
S44 S4 ODS GOO, ESatIIVUSEES49—35)I)
S5S6 sitharamii S50; 351; spicifer
350, 351; sporoboli 342; tripogonis
OS), oy), GNA sik, S58, seyh, sSe tiei=
ie) Sis Wievoriae GO, sei, see

Codinaea 419, 425, 426; apicalis 426;
apiculata 425, 450; aristata 426; as-

samica 426; botulispora 426; brevisetu-
la 425; britannica 426, 450; clavulata
426, 451; cylindrospora 426, 450; di-
morpha 451, 453-455; eucalypti 425,
450; fertilis 426, 455; glauco-nigra
426; gonytrichodes 426; hughesii 426,
450; illinoensis 419, 420, 425; longi-
spora 426; lunata 426, 450; lIunulospo-
ra 421, 422, 425; maharashtrensis 450;
matsushimae 423-425; novae-guineensis

425, 450; obesispora 425; pakhalensis
426, 451; parva 426; septata 450; se-
tosa 426; simplex 426; unisetula 425,
450; vulgaris 426

Coelocaulon see 86-101

Coelomomyces milkoi 232, 482, 483, 494

Coenogonium see 86-101

Collema see 86-101

Coltricia 183; sideroides 183

Completoria 200, 210, 481, 483-487, 490,
492, 494, 502; complens 482, 484-486,
488, 4905 9494, 5503

Conidiobolus 192, 196, 199, 201, 203-
DOS meet =200 su 2iie ae 2i2, G21, Z2h7. 2219,
22 2 2 2G AEN re Oe, EO) ou aOe,
AO Om CU OO Rm Ao 2pm Oo oO OP OOOO 2.
adiaeritus 204; apiculatus 231; corona-
iws AO, WSs imaler Als Ooscurus Aly,
22S MCC eS OAD Abuse > lneatinicOm—
sorceSs 4B, BOO, Bie, Al, Bs 2A.
Zo lpm Al, O12 13

Coniothyrium 371
Conotrema see 86-101
Coprinopsis friesii 376; phaeosporus 394

Coprinus:374, 386, 390, 401; sect Herbi-
COlaeua3 73. 374, oor C90, 392, , 400,
AmMoNiIwS B/S, SM, BO, s/s eirceMue=
ss /4050 380. 98000, 3990;,1295, .990% au
strofriesii 373, 385, 378,. 392, 394,
395; brassicae 373, 378, 386, 388,
39069 394.0 3972 DUrKIIg S15, 5925) ClInCho=
Nensis, 3765, 38858392, (395; friesit 373,
S75 ee OMNES (On msO LO) oC Os Ie,
393, 398, 400, 404; v microspora 398;
herbivorus-— 375, 378, 388, 392, 395,
404: -herinkii 374, 376, 394, 397; kubi-

ora

Gree S73, SUA, Ss, Sele, ceil, Sear
maysoidisporus 373, 374, 380, 381,
382, 384, 386, 387, 389, 390, 393,
396, 400-402; melo 373, 378, 388, 390,
SE" s microspermus 398; neotropicus
373, '375, 390, 394, 395; paleotropicus
$37 o 55 OD, 1090), 394. ©3952" pallidisporus
0985 phaeosporus 373; 375, 9376.) 386,
Sh, SS, SS, S87, OO, 2O49 w sollitie=
rius 398; picosporus 400; platypus
398; pseudofriesii 373, 376, 394, 396,
S973) | psychromorbiduss 373, 95379, moro,
378, 382, 384, 386, 387, 390-393, 396,
400-402; pusio 400; rhombisporus 373,
SUD, Sil 398, 404s Roecinesi Zoe
ScileiMice 37/5, S73, SH Sse sweiliitzeyei=
us 398; subtigrinellus 376, 380, 395,
3965) “subunticicolal 373, )13/8. 386.1590,
S73 iieirinelius S76, S60, S85, cess
triangulospora SfeVy40 urticicola SHS
SH), S73, S82, BS, cei, S80, aA,
397, 399, 400; willapaensis 381; xan-
INNCMS SHS, SHS, SA, Sa, sey

Cora pavonia 90

Coriolopsis asper 179, 180; brunneo-
leuca 181; caperata 179, 182; polyzona
(Sie sancguinatiag | 7Chun 1S lof estrhu—
mosa 185

Cornicularia see 86-101

Corticium cremeoisabellinum 125; subilla—
queatum 126

Corynetes 363, 365; arenarius 365; atro-
purpureus 364, 365; globosus 365; pur-
purascens 365; robustus 365

Corynophoron colensoi 326

Crocicreas 69

Cullicicoia $86, 19, As, Bil, BAsiy 2s.
486

Curvularia 411

Cyclomyces 181; fuscus 181; setiporus
177; tabacinus 182

Cylindrotrichum 257; curvatum 258; el-

lisii 257; gorii 258; helisciforme 258;
oblongisporum 257; proliferum 257; tri-
septatum 257; zignoéllae 258

Cypheliopsis bolanderi 90

Cyphelium inquinans 90

Cyrenella 268, 275, 276; elegans 267,
268-270, 272-276

Cystocoleus ebeneus 90

Cytospora 371

Dacryopinax 428, 430; dennisii 430; ele-
gans 430; indacocheae 429, 430; mar-
tinii 429, 430; maxidorii 428-430; spa-
thularia 430; yungensis 430

Dactylina see 86-101

Daedalea 176; aulaxina 176; flavida
176; fuliginosa 176; lurida 176; micro-
zona 176; plumbea 176; pruinosa 176;

splendens 176; violacea 176
Darbishirella gracillima 90
Dasyscypha coerulescens v dealbata 148
Dasyscyphus 6, 14
Delacroixia 207

930

Dendrographa see 86-101
Dermatocarpon see 86-101
Desmazieria see 86-101
Dictyochaeta 451; fuegiana 451
Dictyonema see 86-101
Dictyopanus 177

Dimelaena see 86-101
Dimeromyces aberrans 470
Dimorphomyces 470

Diploicea canescens 90
Diploschistes see 86-101

Dirina see 86-101

Dirinaria subconfluens 96
Dolichocarpus chilensis 90
Drechslera 339, 344; tripogonis 342, 344

Embellisia 18, 24

Empusa (41; 229,;_, 513; 914; subg Triplo—
sporium 219; aphidis 514; apiculata
7, Pile AKO calrolinieine we, Als.
227, 512; dipterigena 219; forficulae
512; fresenii 219, 227; geometralis
509; major 201, 211, 216; muscae 227;
occidentalis 509; papillata 211, 216;

thaxteriana 223, 224;
Endocarpon see 86-101
Enterographa see 86-101

virescens 213

EntomopHhaga 192, 196, 199, 201, 209-
7a), alc. ASS. ASH), ES. | AS. §=— BOWE
grylli 199, 221; obscura 224; thaxter-
jana 224

Entomophthora 192, 195, 196, 198-200, 203-
20S ePZO9=2 0, e219 e220, 225, 220 eco,
231-234, 473, 483, 486, 487, 499, 501,
304, SiS, Bille eplyicis 2OA, BWA, AS.
SOST OL, @ SI4ee eapieulata ) 204-ae225.
231; aquatica 213; batkoi 232; bulla-
ta 473; canadensis 509; caroliniana
SIZ ecarpentl er immcO4 see 205 seal ae acle.
Zi Gee e2iiiam OlOs ms coleopterornummrclleam lide
218, 509, 510; creatonotus 474; crusto-
Se 809s euilieis 198, ZOO, Bl@, eile
curvispora 229, 230; delphacis 212;
delpiniana 212; elateridiphaga 509; e-
rupta 211; exitialis 511-513; ferrugi-
nee SA, lds ieriiculles OY, Sis cle
gantea 232; gloeospora 194;) grylli
228, 232; henrici 509, 510; ignobilis
Z\\ 9 2S) 2 ch | © (pe kOe 2
ASS TAUISCES IS. Wet ZO, IO, Avil.
487, 501; nebriae 509, 510; obscura
USE), 7, 7S, 27IND CreciceiMMalis ils
ovispora 229, 230, 508; papillata 204,
225, 231; phytonomi 509; planchonia-
met 70, APS [peKeli@eins 4s, 2742), 10%.
509; sphaerosperma 41, 47, 202, 210,
228, 507, 514; thaxteriana 199, 219,
Jixs PA “ANeltovclithin ZBlile TWHslinaie
yey, Gish, SIS), 6 AO), «= AIG weetrrinie@lle
481, 482, 486, 487, 489, 491, 499; vi-
euler: A> Aol, Ail, AiGs Ae “Pil.
513; weberi 210; zabrii 511

Ephebe lanata 90

Erynia 192, 194, 196-198, 201-205, 208,
ZN 213 ee zlO=2 One 2.0) Gee COZ ZO =a 5 Or
AT, 472s aa GO 480. 499 O02 ee D06—
514; americana 473, 511; aphidis 509,

514; aquatica 213, 511; arrenoctona
203; blunkii 511; brahminae 511; bul-
lata 471, 472-474, 479, 511; callipho-
rae 511; canadensis 509; caroliniana
IG, USS), ZAOZ=70, AG, 2S, Sls, SilAs
castrans 512; coleopterorum 509; coni-
ca 225, 511; crassitunicata 509; crea-
tonoti 471, 474, 511; crustosa 509;
curvispora Slits delphacis 212-215,
511; delpiniana 212-214, 511; dipteri-
gena 511; echinospora 511; elateridi-
phaga 509; erinacea 217, 511; formi-
cae 471, 475, 476, 478, 479, 511; geo-
metralis 509; gloeospora 511; gracilis
511; henrici 509, 510; ithacensis 212,
217; jaczewskii 509; lanceolata 509;
magna 512; montana 511; myrmecopha-
Gel CYAl, EMS. Uy, Mis. Sis meoriac
509; neoaphidis 42, 202, 214-216, 511,
Byi[ dive mouryt =) 511, 513; occidentalis
509; orientalis 509; ovispora 507, 508,
510, 512, 514; phalangicida 512; phal—
loides 509; phytonomi 509; radicans
509, 513, 514; rhizospora 512; sepul-
chralis 512; variabilis 512; virescens
512; vomitoriae 512

Evernia prunastri 90
Everniastrum see 86-101
Exserohilum 339

Favolus 176; brasiliensis 176, 177; fi-
brillosus 176; fissus 176; granulosus
176; guadeloupensis 176; junghuhnii
177; multiplex 177; peltatus 177; phi-
lippinensis 176, 177; spatulatus 177;
tener 177; tenuissimus 177

Fibuloporia subvermispora 109

Filobasidium 271

Fomes fasciatus 183

Fomitopsis 183; rhodophaeus 183; scutel-
lata 183; supina 181, 183

Fulgensia desertorum 87

Fusarium roseum 370; semitectum 371

Galactinia 462; megalosperma 462; pseu-

dosuccosa 461, 462; f macrospora 462
Ganoderma Ad, 182-184; applanatum
182, 184; lucidum 182-184
Geoglossum 361, 365; glutinosum 362; ni-
gritum 362; v nigritum 362; umbratile
v umbratile 362
Globifomes graveolens 178
Gloeocystidiellum 36; furfuraceum 39;
propinquum 36; sibiricum 36
Gloeocystidium ltacticolor 36

Gloeoglossum 361, 362; glutinosum 362,
363

Gloeoporus 177; leptopilus 177; pusillus
lia

Glypholecia scabra 91
Graphis see 86-101
Gymnoderma see 86-101
Gyrostomum scyphuliferum 91

Haddowia 182
Haematomma see 86-101

Halysiomyces 407, 408; saxatilis 408

Haptoglossa 483
Helminthosporium 339
Helocarpon 290; crassipes 291

nelionivuun S85 CO, CA, @s, Ws GOR flllexo=
punctatum 62; carpinacola 69; cauda-
tum 62, 83; conocarpi 62; dearnessii
62; epiphyllum 66, 68, 74, 76; errati-
Clin (9s, We, Gee ieuiclicswinn GE, 740).
72; fraternum 72; immutabile 68-69,
73, 74, 76, 83; linderi 76; midlanden-

SO Wos55, GD, UWS, Hes

80; phyllophilum 79;

translucens 80, 83
Heppia lutosa 91
Herpomyces 470
HERPOMYCETACEAE 469
Herpomyceteae 469
HERPOMYCETINEAE 469
Heterobasidion annosum 183
Heterodea muelleri 91
Heterodermia see 86-101;

phyllogenon 79,
rufo-corneum 59;

barbifera 85,

101; circinalis 85, 101; stellata 85,
102; verrucifera 85, 86, 102

Heteromyces 328

Hexagonia 177; blumei 177; cingulata
177; dregeana 177; glabra 177; hirta
185; molkenboeri 177; pulchella 177;
tabacina 177

Hubbsia lumbricoides 91

Humaria gregaria 5

Hydropus 150, 168, 169

Hydrothyria venosa 91

Hymenochaete 241, 254; acanthophysata
MES). PAM). PI, PY, Ae, HSS Clininetino—
mea 254; digitata 241, 246-249, 254,
256; harpago 241; hauerslevii 241,
253-256 ; pinnatifida 241-246, 254,

256; separabilis 241; spathulata 241

Hymenoscyphus 59, 60, 62, 65, 69, 72-
(1 (9. SOO S ealiDopuinetatusso2.cau—
Cais CO=54, G5, Wa, C8, 74%, Us,

cereus 61; dearnessii 60, 62-64,

epiphyllus 60, 65, 66; erraticus

61, 67, 69; fraternus 72; immutabilis

60, 65, 73, 74; lasiopodius 61; leucop-

61; musicola 61; phyllogenon 69;
phyllophilus 79, 80; rufocorneus 80,
sclerogenus 61; scutulus 61; serotinus
61, 72; translucens 61, 80, 82; umbili-
catus 60

Hyphodiscosia europaea 455

Hyphodontia setulosa 111

Hypocenomyce friesii 92

Hypochniciellum 124, 125; cremeoisabelli-—
num 124, 125; molle 124, 125; ovoide-
um 124, 125; subillaqueatum 124-126

Hypogymnia see 86-101
Hypotrachyna bostrychodes 94

Icmadophila ericetorum 91
Ingaderia pulcherrima 91

Knightiella splachnirima 100
Koerberia biformis 91

Laccaria trullisata 267, 269

Lachnellula

145, 148; pittospori ssp azo-
rica 145, 146; ssp pittospori 146; pul-
veracea 145, 147; viridi-glauca 145,
148

Lachnum 139; aranea 138

Lamia culicis 217
Lamprospora ovalispora 7
Lasallia see 86-101

Laxitextum = 535, G36" sbicolor SS ams6.9s9.

40; incrustatum 35, 37, 39; lutescens
39536, 938-40

Lecanactis see 86-101

Lecanidion 350

Lecanora see 86-101, 103; pinguis 102;
pseudopinguis 85, 102, 103; sulphurea
104; texana 85, 103, 104

Lecidea see 86-101, 291; crassipes 291

Lecidella elaeochroma 93

Leciophysma finmarkicum 447; furfur—
ascens 447

Lenzines vrs ecule WHS, I77, Ue, Webbe
mel Melevi uve lesvuiliine: 177, vee ceilt=
ata 178; elegans 178; guilleminiana
178; junghuhnii 178; juvenile 178; mu-
rina 178; myriophylla 178; platyphyl-
las ee platy podalsl(Seaestemtisms izes
vespacea 176-178

Leprocaulon see 86-101

Leptogium see 86-101

Leptosphaeria 350

Leptosporomyces 124; ovoideus 124, 125

Letharia see 86-101

Leucogyrophana 124; cremeoisabellina
124; mollis 124; subillaqueata 124

Lichen cereolus 302
Lichina see 86-101
Lignosus sacer 183
Lobaria see 86-101
Lopadium pezizoideum- 94

Macrobiotophthora 481, 486, 498, 499,
501, 502; vermicola 484, 485, 488, 490,
499, 501-503; vimariensis 482, 484,
485, 487-490, 498, 503

Macrosporium longipes 28

Marasmius 150, 168; decurrens 152, 153,
(56, eee ineasiinesws WSi0),, Yo4%, ies y
CangdiiGissiimusml 2pm OsmE Vann VeUSmtoZ.
15350 IS6serhododencipia io Ipmlos

Maronella laricina 94

Masonhalea richardsonii 88

Massospora 50, 51, 53-55, 198, 200, 205,
206, 210, 211, 227, 486, 492, 501, 502;

cicadettae 50, 51-53, 56;
cicadimal 51) 535 85am cZilteachticeroproc—
tae 56; diminuta 55; dorisianae 53,
Bas wielhiiintee Sse llewiSjevels SiO), Sih

carinetae 55;

55; ocypetes 55; pahariae 50, 54-56;
platypediae 56; spinosa 55; tettigatis
53855

Mastodia tessellata 94

Melanaria melanospora 94

Melanelia see 86-101

Menegazzia see 86-101

Meristacrum 192, 200, 204, 208, 210,
2320 481, 483, 485, 486, 494,

Oe!

[Meristacrum] 495, 498, 502; asterosper-
mum 482-491, 494, 495,, 501, 503;
milkoi 232, 484, 485, 488, 490, 494,
495, | 503; pendulatum, 481-483, 485—
487, 489, 491, 495

Micarea denigrata 94

Microglossum 361, 363, 365; atropurpure-
um 364; olivaceum 363, 364

Microporus 182; affinis 182; scopulosus
182

Microthelia micula 94

Miyoshia fusispora 257

Miyoshiella fusispora 257

Moellerodiscus 62, 79

Monoicomyces leptochiri 416

Monosporium minutissimum 371

Mortierella alpina 370; gracilis 370;
bellina 370; nana 370; vinacea 370

Mucor fragilis 370; varians 370

Muiogone 207

Mycena_ 150, 168,
150, 156; rorida 168

Mycoglaena myricae 94

isa-

IGS: kalalochensis

Nematophthora gymnophila 483
Neofuscelia see 86-101
Neophyllis melacarpa 94
Neozygites 196, 198, 208, 219,
dis 219; turbinata 194
Nephroma see 86-101
Nephromopsis see 86-101
Neuropogon see 86-101
Nia 276
Nigroporus durus 183; vinosus 184
Normandina pulchella 94

228; aphi-

Ocellularia alba 94
Ochrolechia see 86-101
Omphalaria kansana 94
Omphalia rhododendri 150,
Omphalodium arizonicum 91
Omphalopsis rhododendri 152
Opegrapha saxicola 94
Osoriomyces 412, 416-418;
UWB WA\Sy We AT

Oxyporus cervino-gilvus 185

152

rhizophorus

Pachyella 367, 457, 458, 463; adnata
458, 463; aquatilis 458; babingtonii
368, 458, 463; clypeata 458, 460, 463;
hydrophila 457, 458, 463; megalosper-
ma 459, 463; peltata 457, 459, 460,
463; pseudosuccosa 457-459, 461, 463;
punctispora 457, 458, 461, 463; viola-
ceonigra 462, 463

Panellus 177; pusillus 176

Pannaria see 86-101, 434, 445; pezizoi-
des 434; rubiginosa 445

Paracnhnopeziza miniopsis 14
Parathelium see 86-101
Parmelia see 86-101
Parmelina galbina 95
Parmeliopsis see 86-101
Parmotrema see 86-101
Peccania see 86-101

Peltigera see 86-101
Peltula see 86-101

Penicillium 370, 371; asperum 371; chry-
sogenum 370; coryophilum 371; freque-
tans 370; funiculosum 370; kKapuscin-—
skii 371; pinetorum 370; restrictum
370; solitum 370; soppi 370; thomii
370

Pertusaria see 86-101

RPezizal 45/7, 4585) ~aquati lism 457.5) 460i,
462, 464; atroviolacea 368; candido-
fulva 13; caudata 62; epiphylla 66,
74, 76; exidiiformis 458; leonina 13;

phyllophila 79, 80;
violaceonigra 462

Pezoloma laricina 14

Phaeographina see 86-101

Phaeographis see 86-101

Phaeotrichoconis 338;
338; crotalariae 338; urariae 338

Phellinus 178-180; appositus 178; calli-
morphus 179; chryseus 179; extensus
180; fastuosus 180; gilvus 179, 184;
pectinatus 181; senex 178, 181

Phialea dearnessii 62, 65

Phlebia subochracea 111;

Phylliscum see 86-101

Phylloporia chrysita 184

Physcia see 86-101; barbifera 101

Physconia see 86-101

Physma byrsinum 97

Pilophoron 290; sect Eupilophoron 316,
317; sect Nigricaulia 316, 317; cereo-
lus 315; polycarpum 317; robustum 97

Pilophorus 289-293, 295, 316, 317, 326,
328; sect Eupilophorus 291; sect Nigri-
Cauilew 29S acictilainiisu 292.2 o Sc Or
SO), S05, S07, 409), SI@, s/, S20, Se5s
v conjugens 299; f hallii 308; awas-—
thianum 292-298, 300-302, 316; cario-
SUM S495 CGareoliUis S10, 7274, 7885, 2S.
297, 302-307, 31355320," 324; Vo cephalo—
diferus 305; v_ hallii 308; clavatus
292-298, 308-310, 316, 317; colensoi
295, 326; v reagens 326; conglomera-
JU Zk, C86, 329, seis euriullum. 2O2Z;
M28), Vise), PSO, COA, SNO=SUF, SMA, Siles
Cistems ss Wilowile 287, 288, 286)
PNG, 7A}. SIVA, S07, SUES. S28 inal=
fii 308-310; japonicum 308, 310; nigri-
caulle 2912297. 30419 9315-3170 325, 3205
pileatum 328; robustus 290, 292-294,
296-3005 9306-8 1317-32 Ihe 328-9252 Can
distans 321; f magnus 320; staufferi
292,295, 93283) strumaticus 292-2947
296-298, 304-307, 313, 316; 320-324:
vegae 292-298, 300, 304, 316, 324-326

Pithomyces 465; funiculosa 465; pavgii
465-468

Placynthium nigrum 97

Platismatia see 86-101

Plectania sect Plicosporae 6, 8;
oides 6, 8

Pleospora infectoria 26

Poculum 73

Polychidium muscicola 97

punctispora 461;

aurata 331, 335-

subserialis 111

cyttari-

Polydesmia 143; fructicola 143; pruinosa
143, 144

Polyporus 178; abnormis 178; aculeatus
178; albomarginatus 178; anisopilus
178; apalus 178; appositus 178; aty-
pus 178; auriculaeformis 178; blumei
Wee bonplandensis 73) botryoides
(Sse DRacy pUSmalicomecal limonpinusml) 29)
callochrous 179; candicans 179; candi-
dulus 179; chryseus 179; cineraceus
179; cinerascens 179; cohaerens 179;
confertus 179; connexus 179; convolu-
tUS sl 9e ecOomiaceus a 79-5 corprugatus
179; cyathiformis 179; demidoffii 179;
dermatodes 180; dilitatus 180; discifor-
mis 180; dissectus 180; dozyanus 180;
elatus 180; extensus 180; fastuosus
180; flabelliformis 182; fuscellus 180;
fuscus 180; gaudichaudii 181; gaya-
nus 181; gibberulosus 181; gossypinus
ined] guadeloupensis Sits haskarlii
181; hasseltii 181; heteromorphus 181;
hymenius 181; inquinatus 181; kickxi-
anus 181; Korthalsii 181; lenis 181;
lenziteus 181; lteucomelas 181; lindi-
gii 181;longipes 182; macropus 182;
mangiferae 182; manubriatus 182; mar-
chionicus 182; mastoporus 182; mega-
loma 182; melanaleucus 182; melaneus
Iss finikelreevcelluis: iteV45 (iiliteleralketine: — iitsvae
moritzianus 182; murinus 182; nephel-
odes 182, 183; nordmannii 183; noto-
pus 183; ostreatus 183; pala 183; pec-
tunculus 183; perpusillus 183; phaeus
183; placopus 183; platypilus 183;
plumbeus 183; rhodophaeus 183; rigi-
dus 183; rudis 183; rugulosus 183;
sanguinaria 178; sclerodermus’7 183;
scleropodius 183; sericellus 183; seti-
porus 180; sideroides 183, 184; sordi-
dus 184; splendens 184; spurcus 184;
stevenii 184; subflavus 184; swartzi-
anus 184; tegularis 184; tenax 184;
tenuissimus 184; testaceus 184; tracho-
des 184; tricolor 184; trigonis 184;
tristis 184; unguiformis 184; vulnera-
tus 184; zollingerianus 184

Poria 107, 111; albipellucida 105, 108;
cinerascens 105-108, 110; lindbladii
108; quercuum 109; rivulosa 105, 107-
110; subvermispora 105, 107-110

Porina see 86-101

Protoblastenia russula 92

Psathyra urticicola 388

Pseudephebe pubescens 86

Pseudocochliobolus 350, 357

Pseudocoelomomyces milkoi 494

Pseudocyphellaria see 86-101

Pseudoparmelia see 86-101

Pseudoplectania nigrella 8

Psilocybe 346, 347; cubensis 346; subcu-
bensis 346

Psilopezia 367; mummularia 367, 368;
nummularialis 367, 368, 459

Psora see 86-101; cerebriformis 85, 104

Doo

Psoroma see 86-101, 433, 434, 439, 44],
445; bryantii 433-435, 448; cinnamome-
um 433, 435, 439, 443, 445-448; foll-
mannii 433, 434, 436, 448; hirsutulum
435; hypnorum 433-435, 445, 447, 448;
paleaceum 435, 436, 447; rubromargi-
natum 435; tenue 433-436, 437, 446,
448; v borealis 433, 434, 439, 441,
443, 445, 447, 448; v tenue 433, 436,
437, 439, 441, 443, 445, 447, 448

Psorotichia see 86-101

Psorula rufonigra 93

Ptychoverpa bohemica 8

Pulvinula globifera 9; niveo-alba 9

Pyrenophora 339, 350

Pyrenotrichum splitgerberi 97

’ Pyrenula see 86-101

Pyrofomes 178, 179; albomarginatus 178;
demidoffii 179

Pyxine pringlei 97

Ramalina see 86-101

Ramalodium succulentum 98

Reinkella see 86-101

Resinomycena 150, 151, 168, 169; acadi-
GmsiS I5, Wai, Wl, WG2, We7s levine
mescens 150, 151, 163, 165, 168; kala-
Iannis ISA, WES, ley, IS, el,
JO45 fuleliiete: WSO, We, W774, IE@=aiG7,
l67eeirhododendriy iS 152. n 5S o>—
lav, Sil, 162, ies

Rhizocarpon macrosporum 98
Rhizoplaca see 86-101
Rhodosporidium 273;
Rhodotorula 267,

dacryoidum 273
275, 276; aurantiaca

DZ See (CO Malactosa2 co
Rigidoporus fusco-lineatus 183; micropo-
rus 184

Rinodina see 86-101

Roccella see 86-101
Roccellaria see 86-101
Roccellina see 86-101
Ruhlandiella berolinensis 5, 9

Saccharomyces cerevisiae 275

Sarcogyne see 86-101

Sarcoleotia 365; globosa 6, 15
Sarcosoma 6; cyttarioides 5, 8
Scelophoromyces 412, 418; osorianus 412
Schismatomma cupressum 99
Schistophoron tenue 99

Schizopelte californica 99

Sclerotinia 405; bresadolae 405, 406;
candolleana 405

Scutellinia 2, 10; erinaceus 10; pennsy!-
vanica 10; scutellata 6, 10; Uumbro-
rum 11

Scytinostromella 39; humifaciens 39

Septobasidium 207

Sepultaria 11; gregaria 6, 11

Setosphaeria 339, 350

Sinotermitomyces 171, 173; carnosus 172,
USA CEBU TWA, WES

Siphula see 86-101

Sistotrema 185; ochroleucum 185

534

Skelophoromyces 418

Speerschneidera euploca 99

Sphaeronaema spinella 370

Sphaerophorus see 86-101

Sporastatia testudinea 99

Sporormia 350

Squamarina lentigera: 92

Stachybotrys atra 370

Staurothele clopima 99

Stemphylium 411

Stereocaulon see 86-101, 289, 292, 307,
326; colensoi 326; fibula 312, 313; pi-
leatum 307

Stereum australe 254

Sticta see 86-101

Stigmatomyces baeri 417

Stomiopeltis 350

Streptomyces 370

Striatosphaeria codinaeophora 451

Strigula elegans 99

Strongwellsea 192, 194, 196, 197, 202-
2, POS, Dl, Bit3, 222, db, ZO,
50254 006, 95061 5125 Castranse 202088216.
Sis tiene WA, ZO, ils, BAS, Se

Tabanomyces 192, 200; 204, 208, 210)
oY. (Nee Hels ii likol 237, 48), Bes.
487, 491, 494

VaiieoiUin Ate ZIE2
zewskii 509, 511

Teloschistes see 86-101

Termitariopsis 207

Hetamiltomyceses | ClewmlkcZ

Tnamnolia see 86-101

Thelephora mollis 125

Thelidea see 86-101

Theloma see 86-101

Thelotrema diminitum 100

Thrombium epigaeum 100

Thuemenidium 361, 363-365;
um 364, 365

Thysanothecium 328; hookeri 100

Toninia see 86-101

Torula bantiana 127, 128, 130, 132

auxiliare 483; jac-

atropurpure-

5 ES

Trametes 185; acuta 185; cervina 179;
crassa 185; cubensis 183; dermatodes
180, 185; hirsuta 179; incana- 185; ma-
rianna 179, 181, 182; membranaceus
178; menziesii 178-182; modesta 178,
182-184; perrottetii 185; scabrosa 184;
trichomallus 185; versicolor 183, 184;
villosa 181, 184; vittata 186

Trapeliopsis wallrothii 93

Trichaptum 185; biformis 179, 181; bys-
sogenus 184; perrottetii 175

Trichocladium 465; pavgii 465

Trichoderma viride 370

Trichoglossum 361, 366; hirsutum v_ hir-
sutum 366

Trichophaea gregaria v intermedia 6,
11; f gregaria 5, 6, 11; f laevispora
Be, 1

Trichophyton 259; mentagrophytes 259,
261, 265; raubitschekii 259, 260-262,
204 ZOD eu eU iin 259 a2 Ollie O49meZ OO

Triplosporium 196, 198, 200, 208, 210
pein, (tH BikS.. ESP

Trypethelium see 86-101

Tubulicrinis 120; angustus 123; cinctoi-
ols A, Ale cliieimc 149, ile Gor
ike) WAGs clidewlleswic 170, i177 25s
hamatus 123; inornatus 123; ovalispo-
PUS 12405 145 128

Tyromyces. 180; caesius 181;
175, 180; floriformis 180

dissectus

Ulocladium 18
18

Umbilicaria see 86-101

Underwoodia beatonii 6, 12

Urnula craterium 12

Usnea see 86-101

, 20, 24; chlamydosporum

Verrucaria laevata 101
Wynnea americana 12

Xanthoparmelia see 86-101
Xanthopsora texana 93
Xanthoria see 86-101
Xylographa see 86-101

/Agoyainiinera Wil, 4, IS, Wee, We. 197’.
AAV =A08,. AVS, Olle 7A, All, BilZ, Bue.
Js), PISA, (MA BOSS), SUG, Sil
subg Erynia 213, 214; subg Zoophtho-
ra 196, 208, 510; aphidis 514; bulla-
ta 473; crassitunicata 201, 202, 212,
509; creatonoti 474; erinacea 213; exi-
tialis 42, 512; ferruginea 512; forficu—
lae 512; jaczewskii 510; lanceolata
212, 509; myrmecophaga 476, 477; occi-
dentalis 41-47; orientalis 212, 509;
oxsweimin AAilPas Yoyayauliieyitetess, CA, Yay (AP).
S025 PACKERS 2/, AWA, ZI@, 22, 22).
506

Zygnemomyces 481, 486, 489, 492, 495,
498, 501, 502; echinulatus 482, 484-
490, 495, 496, 498, 503; pendulatus
484, 485, 488, 490, 495, 497, 498, 503

Zygorhynchus heterogamus 371

PUBLICATION DATES FOR MYCOTAXON

Volume “12, "No. Z
Volume 13, No. 1
Volume 13, No. 2

March= 17," 1981
May 11, 1981
USUPZ er AEN

REVIEWERS

The Co-Editors express their appreciation to the following individu-
als who have reviewed one or more of the papers in this volume
prior to acceptance for publication

ee (Wedge) Rien OLEBERTSON T= Ae MOORE
J. AMMIRATI Je. GINNS G. MORGAN-JONES
R. J. BANDON! Ino Aig LEGO T. NASH
Pee a OA TRA Joe heetiA LUNES DUE eGEER
R. K. BENJAMIN A. HENSSEN Dz W.) ROBERTS
eae GEN 7 Bey R. A. HUMBER Ora em ROGERS
Hamed ee UROSALL SSRs J. W. KIMBROUGH M. A. SHERWOOD
J. W. CARMICHAEL R. F. LANGDON Ro rey SLOP Sin
Jee RAME FR G. A. LAURSEN W. J. SUNDBERG
J. DREW-SMITH F. F. LOMBARD [Peel aA ARES
F.-E. ECKBLAD D. M. MacLEOD J. Wo THOMSON
J. ERILKSSON D. MALLOCH O. VERONA
S. FRANCIS Dan Ee MiCC ADE R. WATLING
W. GAMS M. R. McGINNIS Ne Mates ANE DIET

ERRATA, VOLUME TEN

Page 471, line 12: for 1094: La read 1094: La Palma.

ERRATA, VOLUME TWELVE

Page 280, line 8: for Rossman, J. Wy- read Rossman, L. J. Spielman, J. Wy-

536

Page

182

203

216

217,

46]

line

Leg.

14,

MA.

ERRATA, VOLUME THIRTEEN

> for gal read Gal

: for (Remaudiére) Batko, read Remaud. & Henneb.,
: for form read from

: for M. etcadina read Massospora cteadtna

: for tmnttabults read tmmutabtlts

: for tmmuttbtle read tmmutabtlis

: for H. cauda- read Hymenoscyphus cauda-

: for Hymenoscyphus fastidtosus read Helotium fasttdtosum
: for sporen read spore

tetnseererg. PEMLOCALL LY mabrd cade

: for 4 read 5

7 tOr Poephi lipinens Usereadur aunt iippinensis
: for brasilensis read brasiliensis

“yor Po phil bipinens ts sready? philippinensis
: for brasilensis read brasiliensis

: for berkleyi read berkeleyi

: for Phillipines read Philippines

: for tenius read tenuis

: for tabcinus read tabacinus

: for supinus read supina

: for Phillipines read Philippines

: for Arendholtz read Arendholz

: delete nuclei of

: for EZ. read Entomophthora

: for Z. read Erynta

: for FE. caolintana and E. carpenttert

read Erynta carolintana and Entomophthora carpentiert

EO
LOr
‘for
:V for
Tor
- for
27for
SEeOr
for
27 for
eo
efor
etTor
2 for
SD idaye
LOT
POT.
Sh op i
for

E. read Empusa

E. read Entomophthora

E. read Entomophthora
segregated read segregates

E. read Entomophthora

C. read Contdtobolus
Maraconesian read Macaronesian
Fusareum read Fusartum
Steptomyces read Streptomyces
aspernum read asperum
dengleyae read dingleyae

P. read Psoroma

P, read Psoroma

C. read Codtnaea

parkhalen- read pakhalen-

C. read Codtnaea

aquttilis read aquatilts

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CO-~EDITORS OF MYCOTAXON

G. L. HENNEBERT RICHARD P. KORF
FRENCH LANGUAGE EDITOR ENGLISH LANGUAGE EDITOR
& BOOK REVIEW EDITOR & MANAGING EDITOR
UCL, Place Croix du Sud 3 PlUs Box 264
B-1348 Louvain-la-Neuve, Belgium Ithaca, NY 14850, USA

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be obtained.

EDITORIAL SERVICES AND INFORMATION FOR PROSPECTIVE AUTHORS

Authors prepare their own camera-ready copy after having received comments
from pre~-submission reviewers. Detailed Instructions to Authors. appeared in
MYCOTAXON 1: 3-12. 1974, and 6: 370. 1977. A copy of each will be sent
upon request to one of the Co-Editors.

We are able to provide prospective authors with two aids to publication.
Both are sold at our cost, and are shipped postpaid from MYCOTAXON, LTD.,
P.O. Box 264, Ithaca, NY 14850 USA.

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